Peerj 3954

A massive update of non-indigenous
species records in Mediterranean marinas

Aylin Ulman1,2,3, Jasmine Ferrario1, Anna Occhpinti-Ambrogi1,
Christos Arvanitidis3, Ada Bandi1, Marco Bertolino4, Cesare Bogi5,
Giorgos Chatzigeorgiou3, Burak Ali Çiçek6, Alan Deidun7, Alfonso
Ramos-Esplá8, Cengiz Koçak9, Maurizio Lorenti10, Gemma Martinez-
Laiz11, Guenda Merlo1, Elisa Princisgh1, Giovanni Scribano1 and
Agnese Marchini1
1
Department of Earth and Environmental Sciences, University of Pavia, Pavia, Italy
2
Laboratoire d’Ecogéochimie des Environnements Benthiques, Université Pierre et Marie-Curie,
Banyuls-sur-Mer, France
3
Institute of Marine Biology, Biotechnology and Aquaculture, Hellenic Center of Marine
Research, Heraklion, Crete, Greece
4
Dipartimento di Scienze della Terra dell’Ambiente e della Vita (DISTAV), Università degli Studi
di Genova, Genova, Italy
5
Gruppo Malacologico Livornese, Livrono, Italy
6
Department of Biological Sciences, Eastern Mediterranean University, Famagusta,
Cyprus
7
Department of Geosciences, University of Malta, Msida, Malta
8
Marine Research Centre (CIMAR), University of Alicante, Alicante, Spain
9
Department of Hydrobiology, Faculty of Fisheries, Ege University, Izmir, Turkey
10
Center of Villa Dohrn-Benthic Ecology, Stazione Zoologica Anton Dohrn, Ischia, Italy
11
Department of Zoology, University of Seville, Seville, Spain
ABSTRACT
The Mediterranean Sea is home to over 2/3 of the world’s charter boat traffic
and hosts an estimated 1.5 million recreational boats. Studies elsewhere have
demonstrated marinas as important hubs for the stepping-stone transfer of
non-indigenous species (NIS), but these unique anthropogenic, and typically
artificial habitats have largely gone overlooked in the Mediterranean as sources of
NIS hot-spots. From April 2015 to November 2016, 34 marinas were sampled
Submitted 9 August 2017 across the following Mediterranean countries: Spain, France, Italy, Malta, Greece,
Accepted 3 October 2017 Turkey and Cyprus to investigate the NIS presence and richness in the specialized
Published 24 October 2017 hard substrate material of these marina habitats. All macroinvertebrate taxa were
Corresponding author collected and identified. Additionally, fouling samples were collected from
Aylin Ulman, aylinh.ulman@unipv.it approximately 600 boat-hulls from 25 of these marinas to determine if boats host
Academic editor diverse NIS not present in the marina. Here, we present data revealing that
Mark Costello
Mediterranean marinas indeed act as major hubs for the transfer of marine NIS,
Additional Information and and we also provide evidence that recreational boats act as effective vectors of
Declarations can be found on
page 38
spread. From this wide-ranging geographical study, we report here numerous new
NIS records at the basin, subregional, country and locality level. At the basin level,
DOI 10.7717/peerj.3954
we report three NIS new to the Mediterranean Sea (Achelia sawayai sensu lato,
Copyright
Aorides longimerus, Cymodoce aff. fuscina), and the re-appearance of two NIS
2017 Ulman et al.
previously known but currently considered extinct in the Mediterranean (Bemlos
Distributed under
Creative Commons CC-BY 4.0
leptocheirus, Saccostrea glomerata). We also compellingly update the distributions
of many NIS in the Mediterranean Sea showing some recent spreading; we provide
details for 11 new subregional records for NIS (Watersipora arcuata, Hydroides
How to cite this article Ulman et al. (2017), A massive update of non-indigenous species records in Mediterranean marinas. PeerJ 5:
e3954; DOI 10.7717/peerj.3954
brachyacantha sensu lato and Saccostrea glomerata now present in the Western
Mediterranean; Symplegma brakenhielmi, Stenothoe georgiana, Spirobranchus
tertaceros sensu lato, Dendostrea folium sensu lato and Parasmittina egyptiaca now
present in the Central Mediterranean, and W. arcuata, Bemlos leptocheirus and
Dyspanopeus sayi in the Eastern Mediterranean). We also report 51 new NIS
country records from recreational marinas: 12 for Malta, 10 for Cyprus, nine
for Greece, six for Spain and France, five for Turkey and three for Italy,
representing 32 species. Finally, we report 20 new NIS records (representing
17 species) found on recreational boat-hulls (mobile habitats), not yet found in
the same marina, or in most cases, even the country. For each new NIS record, their
native origin and global and Mediterranean distributions are provided, along
with details of the new record. Additionally, taxonomic characters used for
identification and photos of the specimens are also provided. These new NIS
records should now be added to the relevant NIS databases compiled by several
entities. Records of uncertain identity are also discussed, to assess the probability
of valid non-indigenous status.
Subjects Biodiversity, Conservation Biology, Ecology, Marine Biology, Taxonomy

Keywords Alien species, Expansion, Distribution, Macroinvertebrates, New records, Pathways,
Recreational boating, Vectors
INTRODUCTION
The seas are being rapidly being tainted by many harmful stressors such as climate
change, overfishing, pollution and non-indigenous species (NIS) (Occhipinti-Ambrogi,
2007). The Mediterranean recreational boating fleet is estimated to contain approximately
1.5 million vessels and hosts over 70% of global charter boating traffic (Cappato, 2011).
It is also the world’s most invaded sea, hosting over 700 NIS (Galil et al., 2017), over
half of which have Indo-Pacific origins and have probably arrived via the Suez Canal
(Galil, Marchini & Occhipinti-Ambrogi, in press). The human-mediated transport of
species across boundaries is dramatically altering the natural distribution of marine
biota, impacting biodiversity as well as human well-being (Carlton, 1989;
Occhipinti-Ambrogi, 2007).
Biological invasions are not only important to understand due to their associated
ecological and economic impacts; but they also provide an opportunity to understand
other important biogeographic processes such as long-distance dispersal, rapid
adaptation and range-expansion processes (Viard, David & Darling, 2016). To properly
assess the bioinvasion process and understand the scale of the associated threats, it is first
necessary to have the most up-to-date information regarding species distributions, which
are used to feed the many databases such as the European Alien Species Information
Network (Katsanevakis et al., 2015), the World Register of Introduced Alien Species
(Pagad et al., 2017) and AquaNIS-Information system on aquatic NIS and cryptogenic
species (Olenin et al., 2014). These databases are highly utilized by scientists and legislators
wishing to assess the breadth of the ecological and socio-economic consequences of
Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 2/59

biological invasions by understanding species’ distributions, measuring trends, and
generating ecological models.
Most records of NIS in the Mediterranean Sea originate from occasional or casual
findings, while only a few monitoring programs thus far have specifically targeted
Mediterranean marine NIS, mainly addressing Marine Protected Areas (MPAs, Mannino
et al., 2017), commercial harbors (López-Legentil et al., 2015), or aquaculture sites
(Verlaque, 2001). Recreational marinas have not yet been systematically surveyed in the
Mediterranean, despite the recent international literature indicating they are important
hubs for new species introduction and secondary spreading events (Acosta & Forrest,
2009; Floerl et al., 2009; Clarke-Murray, Pakhamov & Therriault, 2011; Ashton,
Davidson & Ruiz, 2014). Furthermore, several recent records of marine NIS in the
Mediterranean come from marina habitats (Ros et al., 2013; Marchini, Ferrario &
Minchin, 2015; Maric et al., 2016; Ferrario et al., 2017; Steen et al., 2017), suggesting
that marinas are part of the stepping-stone invasion process.
The definition of NIS adopted here is: “An organism introduced outside its natural
past or present distribution range by direct or indirect human activity (European
Environment Agency, 2012).” This definition implies an anthropogenic-assisted
transport via various pathways, albeit intentional or unintentional. The route that a new
species is transported through to a recipient region is treated as a “pathway.” In the
Mediterranean Sea, in addition to shipping and aquaculture (together considered the
principal pathways of global NIS introductions), the Suez Canal is frequently referenced as
another relevant pathway for the migration of Indo-Pacific species (Galil et al., 2017), and
references therein). Each of these “pathways” can have several “vectors” attributed to
them, which is the means by which they were transported (Minchin et al., 2009; Olenin
et al., 2014). For example, the “shipping” pathway can have the following associated
transport vectors: hull-fouling, ballast water and sea chests. There is a high level of
uncertainty associated with many of these pathways and vectors since it is rather
impossible to prove how a species had been transported, although inferential reasoning
on the locality, and proximity to known hubs for NIS introductions such as major
ports, aquaculture farms or the Suez Canal make it possible to put forth scientifically
sound hypotheses. For this reason, a NIS is often defined as “polyvectic species” sensu
Carlton & Ruiz (2005, see definitions), because it could have been introduced by a certain
combination of pathways or vectors.
This contribution presents new records from the first large-scale survey of
Mediterranean marinas for NIS. From April 2015 to November 2016, 34 marinas were
sampled for NIS across the Mediterranean spanning from Spain, France, Italy, Malta,
Greece, Turkey and Cyprus. Additionally, when permitted, boat-hulls were also inspected
for NIS and their captains interviewed about the boats recent travel history since its
last hull-cleaning to investigate if recreational boats indeed do seed new NIS propagules
to marinas they are visiting, i.e., to verify the role that recreational boating plays as a
vector of spread of NIS. Here, we present new NIS records either for the Mediterranean
basin, sub-region, country or locality. The new records are presented by taxa, with
information on the native origin of the species, their global and Mediterranean

distributions, and details of the present record. Here, new records are provided for
32 macroinvertebrate species in Mediterranean marinas and an additional six species
found on boat-hulls but not in the marina.
MATERIALS AND METHODS

The criteria used for marina selection initially included the sub-region to which they
belong, the number of berths (marina size) and popularity as a tourist locality, and in
addition, the possibility of obtaining permissions and feasibility. The number of visiting
vessels to each marina per annum and staying at least one night was also meant to be used
as a proxy for marina selection, however these data were only available for 20 marinas
(Table S1). A total of 34 marinas were sampled in seven countries, along with a subset of
recreational boat-hulls from 25 of the marinas (Table 1; Fig. 1).
When reporting new sub-regional records for the Mediterranean, the sub-regions included
the following sampled countries: Western Mediterranean (Spain and France); Central
Mediterranean (Italy and Malta); and Eastern Mediterranean (Greece, Turkey, and Cyprus).
Marina substrate sampling

We adopted a rapid assessment protocol (Pedersen et al., 2003; Cohen, 2005; Ashton et al.,
2006) targeting all fouling macroinvertebrate taxa. These “rapid assessments” typically
target a predefined list of species, involve an onsite team of experts, and generally last an
hour. Since our strategy also targeted any unknown invertebrate species for collection, and
generally had one or two collectors at most, we increased the typical sampling time to
approximately eight hours per marina to allow for the careful collection and sorting of all
unknown taxa to ensure no new species could additionally be collected. Samples were
taken from each of the central, outer and inner portions of the marinas to guarantee
representative samples. For the larger marinas (i.e., Port Camargue, Cap d’Agde), the
management authorities provided the use of an inflatable boat with a captain to access
hard-to-reach areas, and to ensure the substrate sampling could be completed in one day.
Photos were taken of the non-indigenous biota using either a SONY RXIII (with a
Nauticam housing) camera, or the Olympus Tough TG-3 or TG-4.
A specialized hand-held rigid net with one sharpened edge was built for the marina
substrate sampling (1 mm mesh size, surface 25 20 cm), which extended to a depth of
1.5 m from the pier to scrape the submerged areas of the pontoons and marina walls over
an area of approximately 0.23 m2. Ladders, tires and buoys were also scraped using a
6.35 cm (diameter) paint scraper or manually. Next, the biota were placed in a plastic tray
and immediately sorted to major taxonomic groups into smaller bottles containing a
75–90% ethanol solution (such as crustaceans, molluscs, polychaetes, etc.) for further
laboratory analysis. The smaller-sized taxa were filtered and collected using a sieve
with a 1 mm mesh size. The only exception were the ascidians, which were immediately
preserved in seawater, then later placed in a freezer for 30–90 min (with care taken not
to freeze the sample), then transferred to a 4% formalin/seawater solution for 48 h for fixing,
and lastly preserved in a 90% ethanol solution; this procedure is necessary for maintaining
some rigidity in the specimens structure, necessary for dissection.

Table 1 List of marinas sampled, with corresponding number, geographical coordinates, sampling dates and boat hulls sampled or not (Y, Yes;
N, No).
Country # Locality name Marina name Latitude and longitude Sampling dates Boats sampled

Spain 1 Alicante Marina de Alicante 38.339 N; 0.4799 E 14/11/2016 N
2 Barcelona One Ocean Port Vell 41.376 N; 2.187 E 22/11/2016 N

France 3 Agde Port Principal du Cap d’Agde 43.281 N; 3.501 E 5–18/06/2015 Y
4 La Grande-Motte Port de la Grande-Motte 43.557 N; 4.082 E 02/11/2016 N

5 Le Grau-du-Roi Port du Plaisance du 43.515 N; 4.132 E 16–28/05/2015 Y
Port Camargue
6 Saint-Tropez Port de Saint-Tropez 43.278 N; 6.637 E 1–30/04/2016 Y

7 Cogolin Marines de Cogolin 43.065 N; 6.586 E 1–30/04/2016 Y
8 Sainte-Maxime Port Privé de Sainte-Maxime 43.307 N; 6.638 E 1–30/04/2016 Y

9 Cannes Cannes Le Vieux Port 43.540 N; 7.032 E 19–28/04/2015 Y
10 Antibes Port Vauban 43.585 N; 7.127 E 1–12/05/2015 Y
11 Villefranche-sur-Mer Port de Villefranche 43.698 N; 7.307 E 22–30/11/2016 N
Italy 12 Lido di Ostia Porto Turistico di Roma 41.737 N; 12.250 E 12–19/07/2015 Y
13 Ischia Island Marina di Casamicciola; 40.748 N; 13.906 E 1–11/08/2015 Y
Marina di Sant’Angelo; 40.695 N; 13.893 E
Porto d’Ischia 40.743 N; 13.939 E
14 Sorrento Porto Turistico Marina 40.629 N; 14.375 E 22–29/07/2015 Y
Piccola Sorrento
15 Palermo Marina Villa Igiea 38.142 N; 13.370 E 26–29/07/2016 Y

16 Palermo Porto La Cala 38.120 N; 13.368 E 2–3/08/2016 N
17 Riposto Porto dell’Etna 37.732 N; 15.208 E 17–28/09/2016 Y

18 Siracusa Porto Grande (Marina Yachting) 37.063 N; 15.284 E 15-16/08/2016 N
19 Marzamemi Marina di Marzamemi 36.733 N; 15.119 E 08/10/2016 N

20 Marina di Ragusa Porto Turistico Marina di Ragusa 36.781 N; 14.546 E 1–7/09/2016 Y
21 Licata Marina di Cala del Sole 37.097 N; 13.943 E 5–10/08/2016 Y

Malta 22 Msida Msida Yacht Marina 35.896 N; 14.493 E 1–8/07/2016 Y
23 Valletta Grand Harbor Marina 35.890 N; 14.523 E 11–18/07/2016 Y

Greece 24 Heraklion Old Venetian Harbor 35.343 N; 25.136 E 1–15/11/2015 Y
25 Agios Nikolaos Agios Nikolaos Marina 35.187 N; 25.136 E 18–25/11/2015 Y

26 Rhodes Mandraki Port 36.449 N; 28.226 E 2–11/06/2016 Y
Turkey 27 Istanbul Setur Kalamış Marinas 40.976 N; 29.039 E 28/08/2015 Y

28 Bodrum Milta Bodrum Marina 37.034 N; 27.425 E 9–11/09/2015 Y

29 Datça Datça Marina 26.722 N; 27.689 E 10/10/2015; N
13/05/2016
30 Marmaris Setur Marmaris Netsel Marina 36.852 N; 28.276 E 14–18/09/2015 Y
31 Fethiye Eçe Marina 36.623 N; 29.101 E 19–24/09/2015 Y

32 Finike Setur Finike Marina 36.294 N; 30.149 E 18–27/05/2016 Y
Cyprus 33 Karpaz Karpaz Gate Marina 35.558 N; 34.232 E 21–27/06/2016 Y

34 Famagusta Famagusta Port 35.123 N; 33.952 E 13–19/06/2016 Y

Figure 1 Marinas sampled in this study. Marina numbers correspond to marinas listed in Table 1.
Full-size  DOI: 10.7717/peerj.3954/fig-1
Boat-hull sampling
A preliminary screening was first completed with boat captains/owners before selection to
ensure that their boat had travelled outside the marina in the past 12 months for a
minimum of one night, so that the vessel posed some risk of spreading NIS. Next, and
only with permission from the boat owners/captains, fouling samples were collected from
the boat-hulls and a short survey was completed with the boat owners/captains on the
vessel’s characteristics, hull-cleaning and painting details, and recent 12 months of travel
history. The fouling samples were collected from the boat-hulls using one of three
approaches (which were dependent on authorizations and feasibility): The first approach
involved inspecting the boat-hulls immediately as they were hoisted from the water at the
carénage (haul-out station) for their maintenance routines (cleaning/painting/repairs).
This approach was mainly used in France: Cannes, Antibes, Marines de Cogolin, Saint
Maxime, Saint Tropez and Cap d’Agde as the sampling season provided the optimal
opportunity to use this approach as these routine maintenance procedures normally occur
before the onset of the tourist season, and was used sporadically in other marinas only
when the opportunity presented itself. The boat-hull including niche areas such as the
propeller, propeller shaft, water vents, rudders and ladders were closely inspected and
fouling samples were collected using a paint scraper and aquarium fishing net wherever
fouling biota were found, and quickly transferred to a bottle containing 90% ethanol.
Photographs were also taken from each boat-hull to crosscheck the results and reduce
likelihood of mistakes. The remaining boat-hulls were sampled via snorkeling or, on a
few occasions, by scuba diving but using the same methods as described above. Care was
taken to ensure the sampling strategy did not release NIS propagules into the marina’s
waters by scraping the samples collected in-water directly into small, finely meshed

aquarium nets. All fouling samples were collected by the first author, with the exception of
the boats in Porto Turistico di Roma which were collected by the ports professional scuba
diver, with careful instructions from the first author on what to collect after reviewing
underwater detailed photos of the hulls.
Taxonomic identification
This study focused on fouling invertebrates; plants and algae were not examined. All
macronivertebrate taxa were collected for identification, and samples requiring expert
identification were sent to appropriate experts.
The preserved specimens were observed under a dissecting microscope and, where
needed, taxonomic slides were prepared and analyzed under an optical microscope.
Photographs of magnified specimens or morphological parts were taken directly from the
microscopes using the Olympus TG-4 camera (i.e., for serpulids and crustaceans), or with
the Tescan Field Emission Scanning Electron Microscope series Mira 3XMU for SEM
pictures, with increasing magnification, at 6–19 mm working distance, using an
accelerating voltage of 10 kV, with graphite metallization and detection by secondary
electrons (i.e., for bryozoans). Bryozoan specimens used for SEM pictures were cleaned
beforehand using a combination of bleach and hydrogen peroxide to remove organic
residues. Ascidians were stained with Masson’s haemalum for dissection.
Some of our records refer to species completely new to the Mediterranean Sea, whose
taxonomic identity has been verified morphologically, but still requiring further genetic
confirmation, since they pertain to taxonomically challenging taxa which have often
revealed complexes of cryptic species. Moreover, a couple of our findings include species
not yet properly described scientifically; thus it is not possible to assign a certain
identification until formal descriptions are completed. These records are discussed in
detail to verify the likeliness of representing introduced populations of NIS. To assign
a NIS status for such species, the Chapman & Carlton (1991) criteria were followed
taking into account factors such as: “appearance in local regions where not found
previously;” “association with human mechanisms of dispersal;” “prevalence or
restrictions to artificial environments;” “insufficient active or passive dispersal capability”
and “exotic evolutionary origin.” Records of species found only on boat-hulls but not
in marinas should only be considered as new NIS country records if certain that the boat
did not leave that country’s waters, since boats represent mobile habitats and are hence
affected by an “uncertain occurrence” (see Marchini, Galil & Occhipinti, 2015).
Non-indigenous species status is dependent on their establishment success in a new
locality, and can be defined as either: not established (a single specimen reported in one or
two localities, rare, uncommon), or established (evidence of a reproducing population in
one or more localities, common or abundant). Additionally, a couple of cases are
presented here for “pseudoindigenous species” (see definitions).
RESULTS
Within the framework of this study, a total of 76 NIS were collectively identified from
34 marinas from the seven countries; however, only new country records and interesting

Figure 2 Number of new NIS records per country by taxonomic group. Sampled countries are shown
with their number of new NIS records found in this study (in brackets), and pie graphs show the new
records represented proportionally by taxa. Full-size  DOI: 10.7717/peerj.3954/fig-2
new locality records are presented here. First, we present the number of new NIS found in
this study per country and by taxa (Fig. 2).
This study revealed three species new to the Mediterranean basin (Achelia sawayai
sensu lato, Aoroides longimerus, and Cymodoce aff. fuscina), 11 new subregional records
(W. arcuata, H. brachyacantha sensu lato and Saccostrea glomerata now present in the
Western Mediterranean; Symplegma brakenhielmi, Stenothoe georgiana, Spirobranchus
tertaceros sensu lato, Dendostrea folium sensu lato and Parasmittina egyptiaca now
present in the Central Mediterranean, and W. arcuata, Bemlos leptocheirus and
Dyspanopeus sayi in the Eastern Mediterranean), for an overall number of 51 new country
records and a few new locality records exhibiting distribution expansions. These new
Mediterranean basin and country records are presented (Table 2) with the corresponding
marina numbers in which they were found from Table 1. Additionally, NIS found on
boat-hulls but not in the respective marina, locality or country, are presented as a warning
signal for future monitoring (Table 3). The numbers of new NIS found per marina are
shown (Table 4), and also the new NIS records are presented by country, specifically
12 for Malta, 10 for Cyprus, nine for Greece, six for Spain and France, five for Turkey
and three for Italy (Table 5). Subsequently, all new NIS records are discussed by species
(first ordered by class and family, and then alphabetically by species, see “New NIS
records: notes on individual species” below). The key taxonomic characters used to
identify these species are accompanied as “Supplementary Data,” along with identification
photos taken of our specimens. Comprehensive reviews of global and Mediterranean
distributions for all NIS listed in Tables 2 and 3 are presented below, along with details on
the new record type and if they were found in the marina, on a boat-hull or both.

Table 2 New NIS record table by species, with numbers in locality corresponding to marina details
from Table 1.
Family Species Country and Marina # Record type
Ascidea Clavelina oblonga Cyprus (#34) *
Clavelina oblonga Turkey (#29) *
Phallusia nigra Cyprus (#33, #34) *
Styela plicata Malta (#22, #23) *
Symplegma brakenhielmi Italy (#15) *, CM

Bryozoa Amathia verticillata Malta (#22, #23) *
Amathia verticillata Cyprus (#34) *
Amathia verticillata Turkey (#28, #30) *
Celleporaria brunnea Spain (#1) *
Celleporaria brunnea France (#4, #5, #6, #8) *
Celleporaria brunnea Malta (#22, #23) *
Celleporaria brunnea Greece (#24) *
Celleporaria vermiformis Greece (#24, #25, #26) *
Celleporaria vermiformis Cyprus (#33, #34) *
Hippopodina sp. A Turkey (#32) *
Parasmittina egyptiaca Turkey (#32) *
Parasmittina egyptiaca Cyprus (#33) *
Tricellaria inopinata France (#3, #5) *
Tricellaria inopinata Greece (#24) *
Watersipora arcuata Spain (#1, #2) *
Watersipora arcuata Malta (#22) *, CM

Watersipora arcuata Turkey (#28, #32) *, EM
Crustacea Ampithoe bizseli Cyprus (#33, #34) *
Aorides longimerus France (#5) **
Bemlos leptocheirus Greece (#24, #25) *, EM

Charybdis (Gonioinfradens) paucidentatus Cyprus (#34) *
Cymodoce cf. fuscina Greece (#24) **
Dyspanopeus sayi Greece (#24) *, EM

Erichthonius cf. pugnax France (#5) *
Ianiropsis serricaudis France (#3, #5) *
Mesanthura cf. romulea Spain (#1) *
Mesanthura cf. romulea Malta (#22) *
Mesanthura cf. romulea Greece (#26) *
Mesanthura cf. romulea Cyprus (#33, #34) *
Paracerceis sculpta Malta (#22, #23) *
Paracerceis sculpta Cyprus (#34) *
Paranthura japonica Spain (#1, #2) *
Paranthura japonica Malta (#23) *
Sphaeroma walkeri Greece (#24) *
Stenothoe georgiana France (#5) *
Stenothoe georgiana Malta (#23) *, CM

(Continued )

Table 2 (continued ).
Mollusca Arcuatula senhousia Spain (#2) *
Dendostrea folium s.l. Malta (#22, #23) *, CM

Polychaeta Hydroides brachyacantha s.l. Spain (#2) *, WM
Hydroides brachyacantha s.l. Greece (#24) *
Hydroides dirampha Malta (#22, #23) *
Hydroides elegans Malta (#22) *
Spirobranchus tetraceros s.l. Italy (#18) *, CM

Porifera Paraleucilla magna Cyprus (#34) *
Achelia sawayai s.l. Malta (#23) **
Pycnogonida Achelia sawayai s.l. Italy (#17, #18) **
Note:
Record type: *New country record, **New Mediterranean record; Letters indicate a new subregional record (WM,
Western Med.; CM, Central Med.; EM, Eastern Med.).
NEW NIS RECORDS: NOTES ON INDIVIDUAL SPECIES

Please note that the numbers used in describing the locality of the new records correspond
to the marinas listed in Table 1.
Class: Ascidiacea
Some ascidians whose likely origin is the Northeast Atlantic (i.e., Clavelina
lepadiformis, Ciona intestinalis, Ascidella aspersa and Botryllus schlosseri) have been
excluded from this study which focuses exclusively on NIS. Genetic studies have shown
that these species include different clades in the Mediterranean, some which can be
considered non-native, and in some cases native (Turon et al., 2003; Perez-Portela et al.,
2013; Bouchemousse, Bishop & Viard, 2016; Nydam, Giesbrecht & Stephenson, 2017). These
cryptogenic species (Carlton, 1996), their origins and status require additional genetic
analyses, which exceeds the breadth of the present study, which is based on morphological
characters.
Family: Ascidiidae
Phallusia nigra Savigny, 1816
Potential native origin: Uncertain, could be from the Red Sea, Indo-Pacific, or Western
Atlantic Ocean.
Distribution: First recorded and described from the Red Sea (Savigny, 1816), then in the
Gulf of Guinea and Angola (Millar, 1965), the Arabian Gulf (Monniot & Monniot, 1997),
the Pacific Ocean (Lambert, 2003), Indian Ocean (Abdul Jaffar, Sivakumar & Tamilselvi,
2009), and the Western Atlantic and Caribbean (Van Name, 1945; Bonnet & Rocha, 2011;
Vandepas et al., 2015).
In the Mediterranean, it has only been reported in the Eastern Mediterranean from
Israel, Lebanon and the Turkish Levantine coast (Çinar et al., 2006; Shenkar, 2008;
Izquierdo-Muñoz, Dı́az-Valdés & Ramos-Esplá, 2009), and most recently from Greece,

Table 3 NIS found on boat hulls but not found in the marina or country.
Ascidiacea Clavelina oblonga Cyprus (#33) L
Bryozoa Amathia verticillata Turkey (#31) L
Celleporaria brunnea France (#3, #7, #9, #10) L
Celleporaria brunnea Greece (#25, #26) *
Tricellaria inopinata Turkey (#27) *
Parasmittina egyptiaca Italy (#21) *, CM

Parasmittina egyptiaca Greece (#25) *
Watersipora arcuata France (#7) *
Crustacea Amphibalanus improvisus France (#5) *
Balanus trigonus Cyprus (#33) *
Cymodoce cf. fuscina Greece (#25) L

Ericthonius cf. pugnax France (#3) L
Paracerceis sculpta Turkey (#31) *
Paradella dianae Italy (#20) L

Paradella dianae Greece (#24) *
Sphaeroma walkeri Greece (#25) L

Stenothoe georgiana France (#3, #10) L
Mollusca Dendostrea folium s.l. Italy (#17) *
Saccostrea glomerata France (#10) *, WM

Polychaeta Hydroides homoceros Cyprus (#33) *
Note:
L = Not previously known from the locality, * = Not previously known from the country. Letters indicate a new
subregional record (WM, Western Med.; CM, Central Med.; EM, Eastern Med.).
specifically from Chalkidiki and Rhodes (Kondilatos, Corsini-Foka & Pancucci-Papadopoulou,

2010; Thessalou-Legaki et al., 2012).
New records: This finding represents the first country record for Cyprus (#33 and #34:
Fig. S1A).
Boat-hull records: Found on one boat-hull moored in Cyprus (#34).
Notes: Although its native origin is uncertain, it is considered a NIS in the Mediterranean
(Çinar et al., 2006; Shenkar, 2008). Vandepas et al. (2015) highlighted some uncertainty
regarding some Phallusia nigra Mediterranean records due to resemblances to the also
dark, native congeneric tunicate Phallusia fumigata (Gruber, 1864), and confirmed
the presence of the introduced Phallusia nigra in the Eastern Mediterranean basin.
For this reason, the morphology of the Phallusia specimens collected from Cyprus were
carefully compared to specimens of the native Phallusia fumigata (found in our own
samples from Port Vell, Barcelona).
Family: Clavelinidae
Clavelina oblonga Herdman, 1880
Native origin: Western Atlantic US coast and Caribbean Sea.

Table 4 Number of NIS per sampled marina, using marina numbers given in Table 1.
#. Marina locality and country # NIS #. Marina locality and country # NIS
1. Alicante, Spain 10 18. Siracusa, Italy 16
2. Barcelona, Spain 11 19. Marzememi, Italy 11
3. Cap d’Agde, France 8 20. Ragusa, Italy 14
4. La Grand-Motte, France 7 21. Licata, Italy 11
5. Port Camargue, France 17 22. Msida, Malta 14
6. Saint-Tropez, France 4 23. Grand Harbor, Malta 13
7. Cogolin, France 6 24. Heraklion, Greece 27
8. Saint-Maxime, France 3 25. Agios Nikolaos, Greece 12
9. Cannes, France 5 26. Rhodes, Greece 16
10. Antibes, France 5 27. Istanbul, Turkey 4
11. Villefranche-sur-Mer, France 2 28. Bodrum, Turkey 12
12. Rome, Italy 9 29. Datça, Turkey 9
13. Ischia, Italy 5 30. Marmaris, Turkey 6
14. Sorrento, Italy 8 31. Fethiye, Turkey 10
15. Villa Igiea, Italy 20 32. Finike, Turkey 14
16. La Cala, Italy 16 33. Karpaz, Cyprus 17
17. Riposto, Italy 13 34. Famagusta, Cyprus 18
Distribution: Its initial record is from Bermuda (Van Name, 1945). It is hypothesized to
be an introduced species to Brazil, first sighted there in 1925 (Rocha, Kremer & Fehlauer-
Ale, 2012). In the Eastern Atlantic, it has been reported as NIS in Cape Verde (Hartmeyer,
1912), Senegal (Pérès, 1951), and the Azores (Monniot & Monniot, 1994). It was described
in the Mediterranean half a century after its initial record as Clavelina phlegraea from
southern Italy and Corsica (Salfi, 1929). It was also found in natural habitats on the
Iberian Coast, about 100 km west of Gibraltar (Ordóñez et al., 2016).
New records: This finding represents a first country record for Turkey (Marina #29) and
Cyprus (#34: Fig. S1B), and two new locality records for mainland France (#5, #7).
Boat-hull records: Found on boat-hulls moored in Cyprus (#33 and #34).
Notes: The species identified earlier as Clavelina phlegraea (Salfi, 1929) in the
Mediterranean was thought to be a native species, but recent genetic analysis confirmed it
as the introduced Clavelina oblonga (Ordóñez et al., 2016). In France, it had only
previously been reported in Corsica, so these new records from the French mainland
indicate its possible expansion along the coast.
Family: Didemnidae
Diplosoma listerianum (Milne-Edwards 1841)
Native origin: Northern Sea.

Distribution: This species was first described from England but is well known from
marinas and harbors worldwide including the Pacific Northwest, Panama, Chile, Japan,
Tahiti, Guam, South Africa and Australia (Rocha & Kremer, 2005; Pérez-Portela et al.,

Table 5 New NIS records by country, with countries ordered from west to east.
Country Species Country Species
Spain Celleporaria brunnea Greece Celleporaria brunnea
Spain Watersipora arcuata Greece Celleporaria vermiformis
Spain Mesanthura cf. romulea Greece Tricellaria inopinata
Spain Paranthura japonica Greece Bemlos leptocheirus
Spain Arcuatula senhousia Greece Cymodoce cf. fuscina
Spain Hydroides brachyacantha s.l. Greece Dyspanopeus sayi
France Celleporaria brunnea Greece Mesanthura cf. romulea
France Tricellaria inopinata Greece Sphaeroma walkeri
France Aorides longimerus Greece Hydroides brachyacantha s.l.
France Erichthonius cf. pugnax Turkey Clavelina oblonga
France Ianiropsis serricaudis Turkey Amathia verticillata
France Stenothoe georgiana Turkey Hippopodina sp. A
Italy Symplegma brakenhielmi Turkey Parasmittina egyptiaca
Italy Spirobranchus tetraceros s.l. Turkey Watersipora arcuata
Italy Achelia sawayai s.l. Cyprus Clavelina oblonga
Malta Styela plicata Cyprus Phallusia nigra
Malta Amathia verticillata Cyprus Amathia verticillata
Malta Celleporaria brunnea Cyprus Celloporaria vermiformis
Malta Watersipora arcuata Cyprus Parasmittina egyptiaca
Malta Mesanthura cf. romulea Cyprus Mesanthura cf. romulea
Malta Paracerceis sculpta Cyprus Ampithoe bizseli
Malta Paranthura japonica Cyprus Charybdis (Gonioinfradens) paucidentatus
Malta Stenothoe georgiana Cyprus Paracerceis sculpta
Malta Dendostrea folium s.l. Cyprus Paraleucilla magna
Malta Hydroides dirampha
Malta Hydroides elegans
Malta Achelia sawayai s.l.
2013). In the Mediterranean, its first record was from Italy in 1975 (Lafargue, 1975),
and is now widespread throughout European and Mediterranean waters (Millar, 1969;
Ramos-Esplá, 1988; Koukouras et al., 1995; Çinar, 2014).
New records: This study presents a new locality record for the Turkish Levantine
Sea/Mediterranean coast (#31, #32). During this study, it was also found in France (#5),
Malta (#23: Fig. S1C), Turkey (#28) and Greece (#24 and #26).
Family: Pyuridae
Microcosmus exasperatus Heller 1878
Potential native origin: Unknown.

This species has a broad global distribution from all continental waters, including
remote localities such as Hawaii and the Mariana Islands, but does not occur in Antarctica
(Nagar & Shenkar, 2016).

In the Mediterranean, it was first reported from south-eastern Tunisia in 1998
(Meliane, 2002; Ramos-Esplá, Izquierdo-Muñoz & Çinar, 2013), then from Lebanon
(Bitar, Ocana & Ramos-Esplá, 2007), Israel (Shenkar, 2008), around the Lebanese coast
in 2009 (Ramos-Esplá, Izquierdo-Muñoz & Çinar, 2013), the Aegean Sea of Turkey
(Ramos-Esplá, Izquierdo-Muñoz & Çinar, 2013), and North-Western Cyprus
(Gewing et al., 2016).
New records: This study presents a new locality record for Turkey (#29) as the
southernmost record for Turkey, and a new locality for Cyprus (#33), illustrating its
ongoing expansion.
Notes: Microcosmus exasperatus and Microcosmus squamiger are both present in the
Mediterranean, however, they do not overlap in distributions: Microcosmus squamiger is
present in the Western and Central Mediterranean whereas Microcosmus exasperatus is
only present in the Eastern Mediterranean (Ramos-Esplá, Izquierdo-Muñoz &
Çinar, 2013). Thus, it has been hypothesized these two species invaded via different
entrances to the basin: Microcosmus squamiger via the Strait of Gibraltar and
Microcosmus exasperatus via the Suez Canal (Turón, Nishikawa & Rius, 2007;
Ramos-Esplá, Izquierdo-Muñoz & Çinar, 2013). Noteworthy is that Microcosmus
exasperatus was not found in late 2014 in Karpaz Marina, Cyprus (#33) by Gewing et al.
(2016) when specifically looking for this species; however, we found it present there
in 2016.
Microcosmus squamiger Michaelsen, 1927
Potential native origin: Australia.

Distribution: Globally, this species is found in the waters of California, South Africa,
Hawaii, and the Western Indian Ocean (Mastrototaro & Dappiano, 2008).
In the Mediterranean, it was first reported from Tunisia in 1967 (Monniot, 1981), and is
now found throughout the Western Mediterranean (Monniot, 1981; Ramos-Esplá, 1988;
Mastrototaro & Dappiano, 2008; Turón, Nishikawa & Rius, 2007) and from the Central
Mediterranean: Taranto, Italy and Grand Harbor, Malta (Izquierdo-Muñoz, Dı́az-Valdés &
Ramos-Esplá, 2009).
New records: This finding represents a new locality for Italy around Sicily (#15, #17, #19,
#20: Fig. S1D). From this study, it was also found in Spain (#2).
Family: Styelidae
Styela plicata (Lesueur, 1823)
Potential native origin: Unknown, cosmopolitan species.

Distribution: This species has been reported worldwide (Harant & Vernières, 1933;
Van Name, 1945; Pérès, 1951; Tokioka, 1963; Ramos-Esplá, 1988). It is considered a
NIS in California (Lambert & Lambert, 2003), Gulf of Mexico (Lambert, 2005),
Brazil (Rocha & Kremer, 2005) and the Mediterranean Sea (Maltagliati et al., 2016).

New records: This finding represents a new country record for Malta (#22 and #23). From
this study, Styela plicata is extremely widespread and was found in all sampled marinas
aside from #6, #9, #11, #29 and #33.
Boat-hull records: Found on boat-hulls moored in the following marinas: France (#3, #5,
#7, #10), Italy (#12, #14: Fig. S1E, #15, #21), Malta (#22), Greece (#24, #26), and Turkey
(#31, #32).
Notes: This is a well-known cosmopolitan hull-fouling species found from many localities
across the Atlantic Ocean from Philadelphia (Van Name, 1945) to Senegal (Pérès, 1951).
Recent genetic analysis suggests that its wide geographic distribution is attributed to many
introductions stemming from human-mediated hull fouling, triggering multiple
introduction events (Barros, Rocha & Pie, 2009). Additionally, most records are from
artificial substrates or harbours, also supporting the hypothesis of an ongoing invasion
(Barros, Rocha & Pie, 2009).
Symplegma brakenhielmi (Michaelson, 1904)
Potential native origin: Unknown

Distribution: It has been found in Australian waters (Kott, 2004), the Pacific Panamanian
coast (Carman et al., 2011), and from the Atlantic in French Guianese waters
(Monniot, 2016). In the Mediterranean, it was reported from Israel in the 1950s (as
Symplegma viride Herdman, 1886), then from Lebanon (Bitar & Kouli-Bitar, 2001;
Bitar, Ocana & Ramos-Esplá, 2007), Israel (Shenkar, 2008) and Turkey (Çinar et al., 2006).
New records: This study presents a new country record for Italy (#15), and a new Central
Mediterranean subregional record. During this study, it was also found in Turkey (#31 and
#32: Fig. S1F).
Boat-hull records: Found on one boat-hull moored in Turkey (#31).
Notes: It is likely that Pérès (1958) is referring to this species under the name Symplegma
viride. Antoniadou, Gerovasileiou & Bailly (2016), in their recent update of ascidians found
in Greek waters, warned of a high-likelihood of a Greek invasion due to its proximity
to the Turkish Levantine coast. This study confirms its spread to the Central
Mediterranean. Soon after this finding in Cyprus from June 2016, it was also reported
from Cyprus in Larnaca Bay in November 2016 by Gerovasileiou et al. (2017).
Bryozoa
Family: Candidae
Tricellaria inopinata D’Hondt & Occhipinti-Ambrogi, 1985
Potential native origin: Indo-Pacific Ocean.

Distribution: It is considered a NIS in New Zealand and cryptogenic elsewhere in
the Pacific, from Japan to Taiwan, Australia and the Northeast Pacific (Dyrynda et al.,
2000). It was also reported from the Northeast Atlantic coasts of Great Britain, Ireland,
Belgium, France, the Netherlands, Spain, Portugal and Germany (Dyrynda et al., 2000;

Arenas et al., 2006; Cook et al., 2013). This species has also been transported via
aquaculture and in association with marine debris stemming from the 2011 Japanese
tsunami which landed in Oregon (Calder et al., 2014).
In the Mediterranean, Tricellaria inopinata was first described in the Lagoon of Venice
in 1982 (D’Hondt & Occhipinti-Ambrogi, 1985) and is considered a NIS in the
Mediterranean Sea because the genus Tricellaria, typical of the Indo-Pacific Ocean, was
previously absent from the basin. After its initial Venetian record, it was reported from
Tunisia (Ben Souissi, Ben Salem & Zaouali, 2006), and from several other Italian localities
(Lodola, Savini & Occhipinti-Ambrogi, 2012; Ferrario et al., 2017).
New records: This finding represents new country records for France (#3 and #5:
Fig. S2A) and Greece (#24).
Boat-hull records: Found on boat-hulls moored in Italy (#14), France (#3 and #5), and
Turkey (#27).
Notes: In Europe, it was found on various types of artificial substrates, e.g., boat-
hulls, ropes, docks and also natural substrates (Dyrynda et al., 2000; De Blauwe &
Faasse, 2001). Generally, Tricellaria inopinata is known to establish successfully in marinas
lacking strong freshwater inputs (Occhipinti-Ambrogi, 1991; Johnson, Winston &
Woollacott, 2012; Cook et al., 2013). If it establishes from boat to marina in Turkey, it
would then present a new country record.
Family: Hippopodinidae
Hippopodina sp. A
Potential native origin: Indo-Pacific Ocean.

Distribution: The species Hippopodina feegeensis (Busk, 1884) from the Indo-Pacific and
the Red Sea, was reported as NIS in the Eastern Mediterranean Sea (Powell, 1969; Morri
et al., 1999; Corsini-Foka et al., 2015). However, Tilbrook (1999) had observed strong
morphological variations within Hippopodina feegeensis colonies from different
geographical regions, and some species were later designated to be new species (Tilbrook,
2006). Particularly, Tilbrook (2006) recognised that the true Hippopodina feegeensis is
restricted to the Philippines Islands, South China Sea and Australia, while two other
Hippopodina spp. were left undescribed (named as Hippopodina “feegeensis,” Holothuria
Bank and Hippopodina “feegeensis,” Ethiopia (sic) in Tilbrook, 2006). The material
presented here is most likely conspecific with the still undescribed Hippopodina sp.
collected by Tilbrook (2006) from Massawa Harbor, Erythraea (K. J. Tilbrook, 2017,
personal communication), and is indicated here as Hippopodina sp. A.
New records: This study presents a new country record for Turkey (#32). It was also found
in Rhodes, Greece (#26: Fig. S2B). Recently, Corsini-Foka et al. (2015) recorded
Hippopodina feegeensis from Mandraki Harbor in Rhodes, in the same locality where it
was also collected during this study (at the Three Windmills wall), and those specimens
will likely be re-assigned to Hipppodina sp. A, after a more comprehensive and detailed
taxonomic comparison is undertaken.

Boat-hull records: Found on two boat-hulls moored in Turkey (#32: Fig. S2C).
Notes: This species is morphologically similar to Hippopodina feegeensis, with only a few
varying characters (see Supplementary Data). Further morphological and genetic
comparisons are necessitated to compare the Mediterranean specimens thus far identified
as Hippopodina feegeensis (Powell, 1969; Morri et al., 1999; Corsini-Foka et al., 2015) with
samples from the Red Sea, which will then lead to a proper taxonomic description for
these Hippopodina samples.
Family: Lepraliellidae
Celleporaria brunnea (Hincks, 1884)
Native origin: Northeast Pacific Ocean.

Distribution: It is widely distributed in the Pacific Ocean (British Columbia, Ecuador,
Gulf of California, Hawaiin Islands, Korea and Panama Canal: see Soule, Soule & Chaney,
1995; Seo & Min, 2009). Recorded as a NIS along the North-eastern Atlantic (Portugal
and France: Canning-Clode, Souto & McCann, 2013; Harmelin, 2014) and Mediterranean
Sea (from Croatia, Italy, Lebanon, Turkey: Koçak, 2007; Harmelin, Bitar &
Zibrowius, 2009; Harmelin, 2014; Lezzi, Pierri & Cardone, 2015; Lodola, Ferrario &
Occhipinti-Ambrogi, 2015; Ferrario et al., 2016; Maric et al., 2016).
New records: These findings represent first country records for Spain (#1), France (#4, #5:
Figs. S2D–S2F, #6, #8), Malta (#22 and #23), and Greece (#24). In Turkey, Celleporaria
brunnea was previously found in Izmir Bay by Koçak (2007), and during this study,
three additional localities are added to its previously known Turkish distribution (#28,
#30 and #31), illustrating its wider expansion along the Turkish south-western and
southern coasts. During this study, it was also present all around Sicily (#15, #16, #17, #19,
#20, #21).
Boat-hull records: Found on boat-hulls moored in France (#3, #7, #9, #10) and Greece
(#25, #26), but was not found from the artificial substrates of those same marinas.
Notes: Many species of the genus Celleporaria are tolerant and opportunistic, and may
exhibit invasive attributes (Dunstan & Johnson, 2004). Celleporaria brunnea was reported as
a fouling organism from different substrates, both natural and artificial (Koçak, 2007;
Canning-Clode, Souto & McCann, 2013; Lezzi, Pierri & Cardone, 2015). Furthermore, it can
be easily spread via hull-fouling, but its introduction via the aquaculture trade cannot be
ruled out, as some of the Mediterranean findings refer to sites in close proximity to shellfish
farms (Lezzi, Pierri & Cardone, 2015; Lodola, Ferrario & Occhipinti-Ambrogi, 2015).
Family: Lepraliellidae
Celleporaria vermiformis (Waters, 1909)
Native origin: Red Sea.

Distribution: Apart from the Red Sea, its distribution is not well known (Vine, 1986;
Ostrovsky et al., 2011). However, it has recently been found in the Gulf of Oman

(Dobretsov, 2015). Its first and only Mediterranean record (prior to our new records listed
below) is from Tripoli, Lebanon (Harmelin, 2014).
New records: This study presents new country records for both Greece (#24, #25, #26)
and Cyprus (#33: Figs. S2G–S2I, and #34).
Boat-hull records: Found on boat-hulls moored in Greece (#25 and #26), and Cyprus
(#33 and #34).
Notes: Since Celleporaria vermiformis was previously recorded from only a single
record from a single site in Lebanon, it was not previously considered as an established
species (Harmelin, 2014). However, these five new locality records presented here now
qualify it as an established NIS in the Mediterranean, and signifies its likely spreading in
the eastern portion of the basin.
Family: Smittinidae
Parasmittina egyptiaca (Waters, 1909)
Native origin: Red Sea and Indo-Pacific Ocean.

Distribution: Parasmittina egyptiaca was reported along the Suez Canal (Hastings, 1927;
Harmelin, Bitar & Zibrowius, 2009), in the Red Sea (Ostrovsky et al., 2011), and from the
Indo-Pacific region (Menon, 1972). In the Mediterranean Sea, it has only been reported
from Lebanon (Harmelin, Bitar & Zibrowius, 2009) and Israel (Sokolover, Taylor &
Ilan, 2016).
New records: This finding represents two new country records for Turkey (#32) and
Cyprus (#33).
Boat-hull records: Found on boat-hulls moored in Greece (#25: Figs. S3A and S3B), and
Italy (#21). The Italian finding presents a new Central Mediterranean record for this
species.
Notes: In our samples, Parasmittina egyptiaca was mostly found growing on
Amphibalanus amphitrite (Darwin, 1854) specimens and oysters. The captain of the
boat hosting this species in Italy explained that his home marina was Finike, Turkey (#32),
and he had just recently travelled from there, through Greece to Sicily. Interestingly,
one could expect many similar examples of new country records for Greece as several
dozens of liveaboard recreational sailboats that used to winter in the Finike marina in
Turkey explained to the first author that since 2014, many had collectively relocated their
vessels to now winter in Agios Nikolaos, Crete (#25). Despite thorough sampling
procedures in Agios Nikolaos, this species was not found present in the marina.
Family: Vesiculariidae
Amathia verticillata (Delle Chiaje, 1822)
Native origin: Caribbean Sea.

Distribution: It has a cosmopolitan distribution from tropical to subtropical regions
in the Atlantic and Indo-Pacific Oceans, the Mediterranean Sea and Macaronesia

(Amat & Tempera, 2009; Wirtz & Canning-Clode, 2009; Minchin, 2012; Ferrario,
Marchini & Lodola, 2014; Marchini, Ferrario & Minchin, 2015).
In the Mediterranean, it was first recorded in the Gulf of Naples (Delle Chiaje, 1822)
and is well-known from the following countries: Algeria, Croatia, Egypt, France, Greece,
Israel, Spain, Syria and Tunisia (Marchini, Ferrario & Minchin, 2015).
New records: This finding represents new country records for Malta (#22, #23),
Turkey (#28 and #30: Fig. S3E) and Cyprus (#34). During this study, it was also found in
Spain (#2), France (#4, #5, #6, #11), the Tyrrhenian coast of Italy (#12 and #14), the Ionian
Sea (around Sicily, #15–21), and Greece (#24 and #26).
Boat-hull records: Found on boat-hulls moored in France (#10), Italy (#12: Fig. S3F, #13,
#14, #15, #17, #20, #21), Malta (#22, #23), Greece (#24–26), Turkey (#28, #30, #31), and
Cyprus (#34).
Notes: It was recently confirmed to originate from the Caribbean (see Galil & Gevili,
2014). Due to its rapid growth rate, it can pose ecological and economic impacts by
forming extensive and resistant colonies on many types of artificial substrates
(Ferrario et al., 2016), and can also facilitate introductions of additional fouling species
(Marchini, Ferrario & Minchin, 2015), such as Caprella scaura, which was found to
be intertwined with it in large abundances in La Grand-Motte, France when we
sampled there.
Family: Watersiporidae
Watersipora arcuata Banta, 1969
Potential native origin: Tropical Eastern Pacific.

Distribution: It is a widespread species distributed from the tropical Pacific such as the
Mexican Pacific coast, California and Hawaii, extending down to Australasia (Wisley,
1958; Skerman, 1960; Banta, 1969; Coles, DeFelice & Eldredge, 1999). In the Mediterranean,
it had only been reported from Porto Santa Margherita Ligure in NW Italy and Porto
Rotondo Marina in Sardinia (Ferrario et al., 2015, 2017).
New records: This finding represents new country records for Spain (#1 and #2),
Malta (#22: Fig. S3I) and Turkey (#28 and #32). This also represents an additional
Italian locality record for Sicily (#18: Figs. S3G and S3H, and #20). Therefore, this
study shows this species is now present in all regions of the Mediterranean,
presenting here two new subregional records for the Western and Eastern
Mediterranean.
Boat-hull records: Found on a boat-hull moored in France (#7), but not found from the
marina substrate.
Notes: In this study, W. arcuata was especially abundant in Siracusa, Sicily. The captain
of the boat in Cogolin, France (#7) hosting this species had recently travelled from
Barcelona, where it was also found in the marina from this study. If it does establish in
France, it would then present a new country record.

Crustacea
Cirripedia
Family: Balanidae
Amphibalanus improvisus (Darwin, 1854)
Potential native origin: Western Atlantic Ocean.

Distribution: It is considered NIS in the Pacific Northwest, and is also present in the Sea
of Japan, New Zealand and northern Europe (Foster & Willan, 1979; Zullo, 1979;
Furman, 1989; Iwasaki, 2006).
In the Mediterranean region, it was first reported from the Black Sea in 1844
(Gomoiu et al., 2002). Next, it was found in the Bosphorus Strait, Turkey (Neu, 1935),
which connects the Black Sea to the Aegean Sea. By the late 1940s it was also reported in
Barcelona (Spain), Catania (Italy), and Alexandria and Abukir (Egypt; Kolosvary, 1949).
Marina records: This finding represents a new locality record for Italy (#12). It was also
found in Turkey (#27).
Boat-hull records: Found on boat-hulls moored in France (#5: Figs. S4A and S4B) and
Turkey (#28).
Notes: If Amphibalanus improvisus happens to establish in Port Camargue marina, this
would then present a new country record for France. The captain of a boat hosting
Amphibalanus improvisus in Port Camargue (France #5) had recently travelled from
Barcelona (where it was recorded long ago), as well as the Balearic Islands. Another
captain from Port Camargue also hosting this species had recently only travelled to the
Balearic Islands, so it is likely that Amphibalanus improvisus is present there. The captain
hosting this species from Bodrum, Turkey (#28) had recently travelled to Istanbul, where
this species has long been present, in addition to travelling through Italy and Greece.
Balanus trigonus Darwin, 1854
Native origin: Indo-Pacific.

Distribution: It was first described from the Pacific Ocean (Darwin, 1854), and has a wide
Indo-Pacific distribution extending to the Red Sea. It is considered NIS in the Atlantic
Ocean and Mediterranean Sea, its first Atlantic record coming from Brazil in the 1860s
(Zullo, 1992). It was introduced to the Atlantic coast of North America around the 1950s
to the 1960s (Moore & McPherson, 1963; Gittings, 1985), and has also been reported from
the Eastern Atlantic from the Azores to South Africa.
Its first Mediterranean record was from the Gulf of Catania, Italy in 1927 (Patane,
1927). It was abundant in the Italian Tyrrhenian, Ionian and Adriatic Seas in the 1960s
(Relini, 1968). It is also reported from Egypt (Ghobashy, 1976), Lebanon (Bitar & Kouli-
Bitar, 2001), Turkey (Koçak, Ergen & Çinar, 1999), Greece (Koukouras & Matsa, 1998),
Croatia (Igic , 2007) and Slovenia (Mavrič et al., 2010).
Boat-hull records: Found on boat-hulls moored in Italy (#14 and #15), Turkey (#32),
Greece (#24 and #26), and Cyprus (#33: Figs. S4C and S4D).

Notes: Although reported on boat-hulls in north-western Europe, it has not established in
that region (Hayward & Ryland, 1995). Relini (1968) questioned a lack of other
Mediterranean records for this species despite its earlier abundance in the Italian Ionian,
Tyrrhenian and Adriatic Seas. In Cyprus, Balanus trigonus has not yet been reported
for the country, and the boat captain in Cyprus hosting this species explained that he
had just travelled along Turkey’s Mediterranean coast and also through Rhodes, Greece
since his last hull-cleaning. If this species establishes itself in Karpaz Marina, Cyprus,
where it was found on boats, it would then present a new country record for Cyprus.
This species can also be transported via both the aquaculture or “Live Fish Food Trade”
pathways due to its custom of gluing itself onto other marine species such as shellfish
and crabs (Zullo, 1992).
Decapoda
Family: Portunidae
Charybdis (Gonioinfradens) paucidentatus (A. Milne-Edwards, 1861)

Distribution: This species has a wide Indo-Pacific distribution, including the Red Sea,
eastern Africa, Australia, New Caledonia, Japan (Poupin, 1994, 1996; Apel & Spiridinov,
1998; Apel, 2001), Madagascar (Crosnier, 1962), the Persian Gulf (Naderloo & Sari, 2007)
and Hawaii (Davie, 1998).
Its first Mediterranean record was in Turkey in 2009 from the Kaş-Kekova specially
protected area from the Turkish Levantine coast (Karhan & Yokeş, 2012). A 2010 record
from Rhodes, Greece provided the second Mediterranean record (Corsini-Foka et al.,
2010), which is only about 140 km from Kaş, Turkey.
New records: This finding represents a new country record for Cyprus (#34: Fig. S5A).
Notes: It may have been introduced to the Eastern Mediterranean via ballast water
(Corsini-Foka et al., 2010).
Dyspanopeus sayi (Smith, 1969)
Native origin: Western Atlantic, from Canada to Florida.

Distribution: It spread from the Western Atlantic to the Northeastern Atlantic and also
to the North Sea: Great Britain, France and Netherlands (Ingle, 1980; Clark, 1986). Its
first Mediterranean record was from the Lagoon of Venice in 1991 (Froglia & Speranza,
1993), then next a little south in the Adriatic Sea in the Po River Delta (Turolla, 1999). In
2009, it was found in a Romanian harbor in the Black Sea (Micu, Niţă & Todorova, 2010),
and in 2010 from the Ebro Delta of the Iberian Peninsula, providing the first Western
Mediterranean record (Schubart, Guerao & Abelló, 2012). In 2011, it was collected from
the central-southern Adriatic Sea lagoon of Varano (Ungaro, Pastorelli & Di Festa, 2012),
and in 2011 it was reported in Mar Piccolo, Gulf of Taranto (Ionian Sea, Kapiris et al.,
2014) another known hotspot for NIS, and then in Lago Fusaro (a brackish lagoon north
of Naples), where it was the most abundant crab (Thessalou-Legaki et al., 2012).

New records: This finding represents a first subregional record for the Eastern
Mediterranean and additionally a new country record for Greece (#24: Figs. S5B and S5C).
It was also found in Sicily (#18) from this study.
Notes: Its first Mediterranean record from Venice is hypothesized to have arrived either
via the ballast water or aquaculture vector (Froglia & Speranza, 1993).
Peracarida—Amphipoda
Family: Ampithoidae
Ampithoe bizseli Özaydinli & Coleman, 2012
Potential native origin: Red Sea and Indian Ocean.

Distribution: To date, the species has only been reported from Tanzania and Turkey
(Izmir Bay, Özaydinli & Coleman, 2012). Its distribution may be much wider than
currently known, but this species could have been misidentified as Ampithoe ramondi
Audouin, 1826, following Schellenberg’s (1928) record of “Ampithoe ramondi” (see notes
below).
New records: This finding represents a new country record for Cyprus (#33 and #34:
Figs. S6A and S6B).
Boat hull records: Found on boat-hulls moored in Cyprus (#33 and #34).
Notes: According to Özaydinli & Coleman (2012), specimens from Tanzania identified as
Ampithoe ramondi by Schellenberg (1928) display ischium lobes identical to Ampithoe
bizseli. For this reason, the native origin of Ampithoe bizseli is hypothesized to be the
Indian Ocean, from where it could have been transferred to the Mediterranean via hull-
fouling. Its current presence in two marinas and also on boat-hulls in those marinas
supports the hypothesis of biofouling as a vector for its wider spread.
Family: Aoridae
Aoroides longimerus Ren & Zheng, 1996
Native origin: Northwest Pacific Ocean.

Distribution: It has been reported from Daya Bay, China (Ren & Zheng, 1996) and from
Osaka and Wakayama, Japan (Ariyama, 2004). It has also been recorded from the
Northeastern French Atlantic coast where it is considered NIS (Gouillieux et al., 2016).
New records: This finding represents a new Mediterranean record (#5: Figs. S6C and
S6D), and a new regional record for the French Mediterranean.
Boat-hull records: Found on boat-hulls in France (#5).
Notes: Port Camargue, France, is situated in close proximity to Thau lagoon, the most
important Mediterranean locality for aquaculture farming of Japanese oysters
(Boudouresque et al., 2011). This information and our new record from boat-hulls
suggests that both aquaculture and shipping are possible vectors of introduction, similarly
to what has been indicated for the French Atlantic record (Gouillieux et al., 2016).

Bemlos leptocheirus (Walker, 1909)
Native origin: Red Sea, Indian Ocean.

Distribution: Aside from early records from its native region: Kenya, Tanzania, South
Africa, Suez Canal (Walker, 1909; Schellenberg, 1928; Sivaprakasam, 1968; Myers, 1975), its
first and only Mediterranean record was from Egyptian coast from Port Said, Alexandria,
and Abu Kir in the early 20th century (Schellenberg, 1928; Bellan-Santini et al., 1998).
However, it was considered to be as absent from the Mediterranean, as it had not been
reported since (Zenetos et al., 2017).
New records: This finding represents a new country record for Greece (#24: Figs. S6E and
S6F, and #25), and confirms its presence and reappearance in the Eastern Mediterranean.
Notes: Previous findings of Bemlos leptocheirus in the Suez Canal and from the
Egyptian Mediterranean coast, near the canals entrance, suggest it to have a “Lessepsian
migrant” vector status (Bellan-Santini et al., 1998), especially since it was also recorded
from buoys and boats (Schellenberg, 1928). Our findings support that it should rather
be assigned to the “biofouling or hull-fouling” vector.
Family: Ischyroceridae
Ericthonius cf. pugnax (Dana 1852)
Potential native origin: Indonesia.

Distribution: Ericthonius pugnax has a wide Indo-Pacific distribution including Australia
(Great Barrier Reef, New Caledonia and New South Wales), Papua New Guinea,
Singapore, Japan, Korea, Malaysia, India, Sri Lanka, Madagascar and Mauritius
(Marchini & Cardeccia, 2017 and references therein). It was reported from New Zealand as
a NIS (Ahyong & Wilkens, 2011). Records of Ericthonius pugnax from South Africa on
mussel rafts (Milne & Griffiths, 2013) may also represent an introduction event. In
the Mediterranean Sea, a record of Ericthonius dydimos from the Adriatic Sea
(Krapp-Schickel, 2013) may refer to this species.
New records: This finding represents a new subregional record for the Western
Mediterranean and a new country record for France (#5: Figs. S6G and S6H).
Boat-hull records: Found on boat-hulls moored in France (#3 and #5).
Notes: An Ericthonius species strikingly similar to Ericthonius pugnax was described by
Krapp-Schickel (2013) from the Lagoon of Venice (and to date, has not been reported from
other localities): Ericthonius didymus. The latter presents a strongly posteriorly lobate
pereopod 5 basis, identical to Ericthonius pugnax. Krapp-Schickel (2013) justifies the
establishment of the new species Ericthonius didymus on the basis of differences in
pereopod 5 postero-distal lobe (in Ericthonius didymus only visible in adult males; in
Ericthonius pugnax, visible in both adult and juvenile males), in shape of gnathopod
2 carpus (bearing two teeth in Ericthonius didymus, versus only one tooth in
Ericthonius pugnax), shape of pereopods 3 and 4 basis, as well as body size. However,
a re-examination of Ericthonius material collected in 2012 from the Lagoon of Venice

(A. Marchini, 2012, private collection), together with a cross-comparison of descriptions
and drawings of both Ericthonius didymus, provided by Krapp-Schickel (2013), and
Ericthonius pugnax, provided by Moore (1988), Just (2009) and Azman & Othman (2013),
shows that the differences pointed out by Krapp-Schickel (2013) may not support
the separation between the two species. With regards to gnathopod 2 carpus,
Azman & Othman (2013) showed that the number of teeth in male Ericthonius pugnax
varies with maturity. Furthermore, we observed some males from Venice having a
single-toothed gnathopod 2 carpus, consistent with the description of Ericthonius
pugnax hyperadult males of Moore (1988). The basis of pereopods 3, 4 is bottle-shaped,
and distally expanded in both species. Furthermore, body length is largely variable
(Ericthonius didymus described from Venice is 4.5 mm; Ericthonius pugnax
described from Australia by Moore (1988) and Just (2009) is 3.0–3.7 mm, from
Malaysia by Azman & Othman (2013) is 3.8 mm, from Japan by Nagata (1965) is up
to 7.5 mm.
Therefore, we hereby suggest that the “endemic” Ericthonius didymus in Venice may be
an introduced population of the Indo-Pacific Ericthonius pugnax, and therefore may be a
pseudoindigenous species (see definitions). However, it is also possible that the global
populations of Ericthonius with a posteriorly lobated pereopod 5 basis represent a
complex of cryptic species. We consider that in this case the hypothesis of the valid
introduced status is supported by the following facts:
1. Ericthonius pugnax has a notably wide distribution in the Indo-Pacific region, which
supports a human-mediated dispersal hypothesis, and is already known as a NIS from
New Zealand (and possibly, South Africa);
2. In the Lagoon of Venice, it has developed populations with high densities (A. Marchini,
personal observation), which is consistent with “invasive” behaviour; and
3. The Lagoon of Venice is a well-known hotspot of introductions, where over 70 NIS have
already been recorded, many with Pacific/Indo-Pacific origins, which were introduced
to Venice via oyster imports (Marchini, Galil & Occhipinti, 2015). The present records
from Cap d’Agde and Port Camargue are both nearby another popular hotspot for
oyster introductions, the Thau lagoon (Boudouresque et al., 2011). This further
supports the hypothesis of introduction from the Indo-Pacific region, with aquaculture
being the main pathway of primary introduction.
Family: Stenothoidae
Stenothoe georgiana Bynum & Fox 1977
Native origin: Western Atlantic.

Distribution: Its first record outside its native range was reported just recently in
2010, in association with fouling communities of offshore fish farms (about 10 km from
shore) in Alicante and Murcia, Spain (Fernández-González & Sánchez-Jerez, 2017). Its
subsequent Mediterranean records were from the Ligurian Sea and from Sardinia, Italy
(Ferrario et al., 2017).

New records: These findings represent new country records for France (#5) and Malta
(#23). This study increases its known Italian distribution by incorporating Sicily (#14:
Figs. S6I and S6J, #15, #18, #21). The Maltese and Sicilian findings from this study
represent a new Central Mediterranean subregional record.
Boat-hull records: Found on boat-hulls moored in France (#3, and #10), and Italy (#14,
#15, #17, and #21).
Notes: Since this species has only very recently been reported in the Mediterranean,
we hypothesize that it may have gone previously overlooked, since it is already present in
at least four countries. It may soon establish in Cap d’Agde Marina or Port Vauban,
Antibes, and this should serve as a warning for future monitoring of those marinas. This
study demonstrates that this species is likely polyvectic (see definitions): in addition to its
likely transfer via aquaculture (Fernández-González & Sánchez-Jerez, 2017), recreational
boating is also facilitating its spread.
Peracarida—Isopoda
Family: Anthuridae
Mesanthura cf. romulea Poore & Lew-Ton, 1986
Potential native origin: Tropical to sub-tropical southern seas.

Distribution: Mesanthura specimens belonging to the same species and sharing major
diagnostic characters with Mesanthura romulea described from Australia (Poore & Lew-
Ton, 1986) were subsequently (2000) collected from the harbors of Salerno and Taranto
(Italy), where they were well established (Lorenti, Dappiano & Gambi, 2009), and also
from Ischia Island (Kroeker et al., 2011). More recently, this species has been reported by
Ferrario et al. (2017) from marinas in Northern Italy (Liguria).
New records: This finding represents new country records for Spain (#1: Fig. S7A), Malta
(#22: Fig. S7A), Greece (#26) and Cyprus (#33 and #34), the latter two records also
confirming the presence of Mesanthura cf. romulea in the Eastern Mediterranean. From
this study, we additionally report specimens from Italy (#14, #15, #16: Fig. S7A, and #18).
Notes: The earliest mention of the presence of the genus Mesanthura in the Mediterranean
region was from Lake Burullus, Egypt (Samaan, Ghobashy & Aboul Ezz, 1989); however,
the record was not supported with taxonomic details and needs confirmation. Castelló
(2017) recently described a new Mesanthura species from both the Lebanese coast and
Cyprus (Mesanthura pacoi, Castelló, 2017), whose females vary from those of the present
species in the dorsal colour pattern and in other subtle morphological features. As
mentioned above, the species found by Lorenti, Dappiano & Gambi (2009) and reported
here is comparable and most probably conspecific (G. Poore, 2017, personal
communication) with Mesanthura romulea described by Poore & Lew-Ton (1986), which is
based only on two specimens collected from Sydney Harbor and Port Stephens, New
South Wales, Australia. No other records of this species have been published.
The fact that the extant description of the Australian Mesanthura romulea lacks a
number of taxonomic characters and is based on only two specimens prevents from

determining if features observed in all Mediterranean specimens lie within the natural
range of morphological variation of the species, or allow for the determination of a
different species.
As long as these cases of taxonomic identity are unsolved, and no new material of
Mesanthura romulea is found from its putative native range, the origin of populations
occurring in the Mediterranean remains obscure. However, the Mediterranean finding of
the present species of Mesanthura shows strong indications of a human-mediated
introduction. Following Chapman & Carlton’s (1991) criteria, the lack of previous records
of the genus Mesanthura on a basin scale (except for the recent discovery of Mesanthura
pacoi from the Levantine Sea), the mentioned occurrences from confined areas such as
lagoons and harbors, the notably poor capabilities of active or passive spreading by
natural means of the genus, and its likely exotic evolutionary origin, cumulatively support
the hypothesis of a human-mediated introduction.
Family: Janiridae
Ianiropsis serricaudis Gurjanova, 1936
Native origin: Sea of Okhotsk to the Sea of Japan.

Distribution: In addition to its native range, it has been reported from the Northeastern
Pacific (from Puget Sound to Monterey Bay), the Northwestern Atlantic (from Maine to
New Jersey) and the Eastern Atlantic and North Sea (England and the Netherlands)
(Hobbs et al., 2015).
Its first Mediterranean record was in 2012 from the Lagoon of Venice (Marchini,
Ferrario & Occhipinti-Ambrogi, 2016a), and soon after from Olbia, Sardinia in 2014
(Marchini, Ferrario & Occhipinti-Ambrogi, 2016b).
New records: This finding represents a new country record for France (#3 and #5:
Fig. S7B).
Notes: In North America, this species is now known as a common fouling species. It was
hypothesized that this species is likely more established along North America and the
European coasts than what is known, but may go undetected due to its minuscule size
(<3 mm) and the taxonomic complexity of the genus (Hobbs et al., 2015). All the
Mediterranean findings (Venice, Olbia, Port Camargue) refer to sites in close proximity to
aquaculture sites.
Family: Paranthuridae
Paranthura japonica Richardson 1909
Native origin: Northwest Pacific Ocean.

Distribution: It was first reported from Muroran, northern Japan and from eastern Russia
(Nunomura, 1977). It was reported as a NIS for San Francisco Bay in 1993, then from
southern California in 2000 (Cohen & Carlton, 1995; Cohen, 2005). Between 2007 and
2010 it was first recorded in European waters from the Bay of Biscay, France, most likely
via the aquaculture vector (Lavesque et al., 2013).

Its first Mediterranean records occurred only recently; between 2010 and 2012 it was
found in numerous localities around Italy: the Lagoon of Venice, La Spezia and Olbia
harbors (Marchini et al., 2014), and Taranto (Lorenti et al., 2015). Next, it was found in La
Grande-Motte, France (Marchini, Ferrario & Minchin, 2015) and then in Tunisia and
Greece (Tempesti, Langeneck & Castelli, 2016).
New records: These findings represent new country records for Spain (#1: Fig. S7C,
and #2) and Malta (#23). Furthermore, Paranthura japonica was found in countries
where it was already reported from, extending its known distribution to new localities in
France (#3, #4 and #9), Italy (#12, #13, #16–21: (#21) Fig. S7C), and Greece (#24 and #26).
These new Sicilian records (#16–21), and Maltese record (#23) show it is already
well-established in the Central Mediterranean.
Boat-hull records: Found on boat-hulls moored in France (#3, #5, #9 and #10), Italy
(#12, #17, #20 and #21) and Greece (#24).
Notes: The current findings dramatically increase the known distribution of Paranthura
japonica, revealing it as one of the most widespread NIS in the Mediterranean Sea.
While the initial findings of Paranthura japonica had suggested an association with
aquaculture transfers, these new records show that it most likely is a polyvectic species
(see definitions) species, which complicates the possibility of reconstructing its invasion
trajectory.
Family: Sphaeromatidae
Cymodoce aff. fuscina Schotte & Kensley, 2005
Native origin: Persian Gulf.

Distribution: Cymodoce fuscina was first described in 2005 from seagrass beds in Saudi
Arabia, the Persian Gulf by Schotte & Kensley (2005). Until now, this isopod had not been
reported outside the Persian Gulf.
New records: This finding represents a new record for the Mediterranean basin, and a new
country record for Greece (#24: Figs. S7D and S7E).
Boat-hull records: Found on boat-hulls moored in Greece (#24 and #25).
Notes: Our specimens show very strong affinity to Cymodoce fuscina from the
Persian Gulf (V. Khalaji-Pirbalouty, 2016, personal communication), and they
certainly differ from all other known Cymodoce species reported in the Mediterranean
Sea in several characters of the pleotelsonic region, while also being similar to other
species described from the Western Indian Ocean (Khalaji-Pirbalouty & Raupach, 2014).
Its association with marina structures and hull-fouling further supports the hypothesis
of a human-mediated introduction, possibly from boats travelling from the Red Sea
through the Suez Canal. However, slight differences between our material and the
original description of Cymodoce fuscina by Schotte & Kensley (2005) should be noted,
for example the pleotelsonic apex of Cymodoce fuscina has the three apical lobes
subequal in length and rounded apically, while in our material the central lobe is
slightly longer than the lateral ones, and ends in a tiny bifid spike. We stress the

fact that not all Indo-Pacific species within this genus may be known (many new
species have been described in the recent decade), and a complex of species is also a
possibility. Therefore, we recommend that genetic analyses should be undertaken to
compare the Mediterranean material with specimens from the native range, to
confirm the identity of these samples from Heraklion, Greece.
Paracerceis sculpta (Holmes 1904)
Native origin: California.

Distribution: This is a widely distributed species naturally found along the North
American Pacific coast from California to Mexico, and has also been reported from
Hawaii, Hong Kong, Australia, Brazil and the Azores (Marchini et al., in press and
references therein).
In the Mediterranean Sea, it was first reported from the Lake of Tunis, Tunisia (Rezig,
1978); and next from several Italian localities (Forniz & Sconfietti, 1983; Forniz &
Maggiore, 1985; Savini et al., 2006; Ferrario et al., 2017), and the Strait of Gibraltar
(Castelló & Carballo, 2001). Most recently, it was reported in Thermaikos and Toroneos
Gulf in Greece (Katsanevakis et al., 2014) and La Grande-Motte in France (Marchini,
Ferrario & Minchin, 2015).
New records: This finding represents new country records for both Malta (#22: Fig. S7F,
and #23) and Cyprus (#34). It was also found in France (#4), Greece (#24 and #26) and
Italy (#13, #15–21).
Boat-hull records: Found on boat-hulls from Sicily (#17, #20 and #21), Greece (#24) and
Turkey (#31).
Notes: This species has often been reported from marinas, indicating that recreational
boating plays an important role in the spread of this global invader. In Fethiye (#31), it was
found on a boat-hull but not in the marina and so far was unknown from Turkey; When
interviewed, the boat captain hosting this species explained he had just travelled from
Rhodes (#26), where it was found in the marina. Attention should be paid to see if it spreads
to the marina in Fethiye, Turkey, where it would then constitute a new country record.
Paradella dianae (Menzies, 1962)
Native origin: Eastern Pacific Ocean.

Distribution: The first description of this species was from the Bay of San Quintin,
Baja California (Menzies, 1962).
Its first Mediterranean record was from Civitavecchia, Italy (Forniz & Maggiore,
1985), followed by a series of findings in Egypt (Atta, 1991), Spain (Castelló & Carballo,
2001), Turkey (Çinar et al., 2006), Cyprus (Kırkım et al., 2010), Libya (Zgozi, Haddoud &
Rough, 2002) and Sardinia, Italy (Ferrario et al., 2017).
New records: This finding represents a new locality record for Sicily, Italy (#15: Fig. S7G),
and an additional record for Turkey from the same locality (Fethiye) it had previously
been reported in (#31).

Boat-hull records: Found on boat-hulls moored in Greece (#24), and Italy (#20).
Notes: This species has not yet been reported in Greece, so this finding on a boat-hull in
Heraklion, Crete, which had only travelled through Greek islands since its last cleaning
alludes to its presence in Greek waters. Interestingly, the boat-hull it was found on in Sicily
had just travelled from Fethiye, Turkey, where it is known from. It is assumed that this
sphaeromatid isopod arrived to the Mediterranean via hull-fouling on vessels from the
Northeast Pacific, its alleged original native range (Galil, Occhipinti-Ambrogi &
Gollasch, 2008).
Sphaeroma walkeri Stebbing 1905
Native origin: Indian Ocean.

Distribution: This species is commonly found in intertidal fouling communities and has
been widely reported from ports in warm and warm-temperate waters worldwide,
including the Pacific coast of North America (Carlton & Iverson, 1981).
Its first Mediterranean record is from Port Said, Egypt in 1924, where it was found on
boat-hulls (Omer-Cooper, 1927). Half a century later (in 1977), it was reported from
Toulon, France (Zibrowius, 1992), then from Turkey (Kocatas, 1978), and Alicante, Spain
in 1981 (Jacobs, 1987). Decades later it was found once again on boat-hulls in Haifa
Harbor, Israel (Galil, Occhipinti-Ambrogi & Gollasch, 2008), and also found to be
well-established in Tunisian harbors and lagoons (Ounifi-Ben Amor, Ben Salem &
Ben Souissi, 2010). In 2010 it was first spotted in Italy in the harbor of La Spezia
(Lodola et al., 2012).
New records: This finding represents a new country record for Greece (#24: Fig. S7H).
It was also found in Turkey (#31).
Boat-hull records: Found in Greece (#24 and #25), and Turkey (#31). This presents a new
locality record for Greece (#25) in addition to the new country record presented above.
Mollusca
Family: Chamidae
Pseudochama cf. corbierei (Jonas 1846)
Native origin: Red Sea, Gulf of Aqaba and Suez Canal.

Distribution: It is considered endemic to the Red Sea and Suez Canal (Barash & Danin,
1972). Its first Mediterranean record is from Greece (Ralli-Tzelepi, 1946), and it has also
been reported from Turkey (Cachia & Mifsud, 2017). The latest record from Malta
represents its first Central Mediterranean record (Cachia & Mifsud, 2017).
New records: One juvenile specimen was found in Italy (#20: Fig. S8F).
Notes: This species was formerly known as Chama corbieri, while Pseudochama cornucopia
(Reeve, 1846) and Pseudochama ruppelli (Reeve, 1847) are both considered common
synonyms. An additional record from Israel (Barash & Danin, 1972) as Chama
cornucopiae Reeve, 1846 was excluded since the record was based on an empty shell. The

present finding in Ragusa, Sicily (Italy) of a single young specimen remains dubious about
its exact determination. Hence, we classify this finding as uncertain since the defining
characters for this species were not yet fully developed in our juvenile specimen and
suggest that the occurrence of Pseudochama corbierei awaits further confirmation before
considering the species introduced to Italy.
Family: Mytilidae
Arcuatula senhousia (Benson 1842)
Native origin: Siberian Peninsula to Indo-Pacific.

Distribution: It has been reported from Great Bitter Lake, the Suez Canal, the Red Sea,
Mauritius, Zanzibar, and several Indo-Pacific and Indian Ocean countries including
Thailand, Malaysia and New Caledonia (Barash & Danin, 1972).
Its first Mediterranean record was from Israel in 1960, and then from Lake Bardawil
on the Egyptian Sinai Peninsula in 1982 (Barash & Danin, 1971). It was also found in
Thau Lagoon, France in 1982, a popular oyster aquaculture locality (Hoenselaar &
Hoenselaar, 1989) and then spread to the surrounding area including the Leucate Lagoon.
Next, it was recorded in Ravenna, the Italian Adriatic coast in 1986 (Lazzari & Rinaldi,
1994). In this century, it was found in the Gulf of Olbia, Tyrrhenian Sea (Savarino &
Turolla, 2000), then in 2001 it was established in the Gulf of Taranto, the Ionian Sea,
from an area involving both mussel aquaculture and intense shipping (Mastrototaro,
Matarrese & D’Onghia, 2003). Next, it was reported again along the Adriatic Italian coast
(Solustri, Morello & Froglia, 2003), and the following year it had dense populations inside
the dams of the Port of Leghorn (Livorno, Italy) (Campani et al., 2004; Margelli et al.
2004). It was also found in Tunisia (Ben Souissi et al., 2005), then, between 2006 and 2009,
in Siracusa’s Porto Grande Marina, Sicily (Brancato & Reitano, 2009). In 2010, it was
found in the Eastern Adriatic from the Neretva River Delta growing on serpulid tubes of
the polychaete Ficopomatus enigmaticus (Fauvel, 1923) (Despalatovic et al., 2013). In
Spain, it was reported from the Ebro River Delta in 2014 (Soriano & Salgado, 2014);
however, that record was based on four empty shells, therefore, its presence in Spain still
awaits confirmation from live specimens.
New records: This finding represents the first confirmed country record for Spain
(#2: Figs. S8B and S8C). It was also found in France (#5 and #9), and Sicily
(#15, #16 and #21).
Family: Ostreidae
Dendostrea folium sensu lato (Linneaus 1758)

Distribution: Its first Mediterranean record is from Iskenderun Bay, Turkey in 1998
(Çeviker, 2001), then from Cyprus (Zenetos, Konstantinou & Konstantinou, 2009), and next
from the Greek islands of Astypalaia, Rhodes and Kastellerizo (Karachle et al., 2016). It has
also recently been reported from Panama (Lohan et al., 2015).

New records: This finding represents a new subregional record for the Central
Mediterranean, and a new country record for Malta (#22 and #23). It was also found in
Greece (#24 and #26), Turkey (#29, #30 and #32) and Cyprus (#33: Fig. S8I), where it was
previously known.
Boat-hull records: Found on boat-hulls moored in Italy (#17), Greece (#26), Turkey
(#31 and #32) and Cyprus (#33).
Notes: If it establishes in the marina in Italy, where it was found on a boat-hull, it would
then present a new country record; the boat which was hosting Dendostrea folium in Italy
had just returned from a long trip back from southern Turkey and the Greek Islands.
Dendostrea frons (Linnaeus, 1758) and Dendostrea folium are very similar species.
Huber (2010) rejects the possible presence of Dendostrea frons in the Mediterranean Sea,
despite many reports of this species there. Based on genetic results, Crocetta et al. (2015)
demonstrated that the Greek and Turkish material belongs to a single, morphologically
highly variable species: Dendostrea folium, most likely representing a complex of species
in need of revision (M. Oliverio, 2017, personal communication).
Saccostrea cf. cucullata (Born 1778)

Distribution: It is found from the Red Sea, East Africa down to South Africa including
Madagascar, and West Africa up to Angola (Branch et al., 2002).
Its first Mediterranean record is from south-eastern Turkey in 1998–1999 from
Erdemli, and later from Yumurtalik and Tasuçu (slightly west and east of Erdemli,
respectively), where it is well-established with large populations (Çevik, Öztürk & Buzzuro,
2001), followed by a record from El-Faham, Egypt (Gofas & Zenetos, 2003). An additional
record from Tunisia remains questionable (Ounifi-Ben Amor et al., 2016).
New records: This finding represents a possible new country record for Greece (#24).
From this study, it was also found in Turkey (#31), presenting the most south-western
record for the country.
Notes: The only specimen collected in Heraklion (Greece) was a juvenile (20 mm) and
the crenulations along the margin (a key identification character) were only partially
visible (Figs. S8G and S8H), but were not well developed as in matured specimens.
Therefore, we regard this finding as uncertain and suggest the occurrence of Saccostrea
cucullata needs further confirmation before officially presenting this as a new country
record in Greece.
Saccostrea glomerata (Gould 1850)
Native origin: Australasia.

Distribution: Its native distribution extends from eastern Australia to New Zealand.
In the Mediterranean, it was intentionally introduced to the Adriatic Sea in 1984 for
aquaculture (Cesari & Pellizzato, 1985), but has not been found there since 1990
(Mizzan, 1998), and is thus currently considered as locally extinct. In 1998 it was reported

in Turkey, which was the first Eastern Mediterranean record (Çevik, Öztürk & Buzzuro,
2001), but this record is considered a case of misidentification with either Saccostrea
cucullata (according to Gofas, 2011) or Dendostrea frons (acccording to Albayrak, 2011),
so this record remains questionable.
Boat-hull records: This species was found on one boat-hull moored in France (#10:
Figs. S8D and S8E), which had only travelled locally around the French Riviera (from Nice
to Golfe-Juan) for the past 1.5 years since its last hull-painting.
New Mediterranean records: This finding confirms its presence in French waters and also
presents a new subregional record for the Western Mediterranean.
Notes: This species was formerly known as Saccostrea commercialis (Iredale & Roughley,
1933), and is distinct from Saccostrea cucullata in terms of DNA 16S sequences
(Lam & Morton, 2006; Salvi, Macali & Mariottini, 2014).
Polychaeta
Family: Serpulidae
H. brachyacantha sensu lato Rioja 1941
Potential native origin: Mexican Pacific.

Distribution: Since its initial Mexican record, it has been reported globally, from
Hawaii (Straughan, 1969), Brazil (Zibrowius, 1970), Micronesia (Imajima, 1982), Japan
(Imajima, 1987), Venezuala (Dı́az Dı́az & Liñero-Arana, 2001), California (Bastida-Zavala
& ten Hove, 2003) and India (Pati, Rao & Balaji, 2015).
Its first Mediterranean record was from Israel in 1933 (Ben-Eliahu, 1991), and its
second from Turkey (Çinar, 2006).
New Mediterranean records: This finding represents new country records for both
Greece (#24) and Spain (#2: Figs. S9A–S9C), the latter also presenting a new subregional
record for the Western Mediterranean.
Boat-hull records: Found on boat-hulls moored in Greece (#24).
Notes: The recent paper by Sun et al. (2016) re-described H. brachyacantha as a complex
of species, which renders the identity of the Mediterranean populations as unknown,
until genetic analyses are performed and the status of the species within the complex is
clarified. Consequently, the native origin of the Mediterranean populations is also
unknown, and this serpulid should therefore be classified as “cryptogenic.” However, the
possibility that H. brachyacantha is a native Mediterranean species having long escaped
detection is not fully supported; it first appeared in the Mediterranean as early as in
1933 and so far has only two records in the Levantine Sea (Israel and Turkey). According
to Chapman & Carlton’s (1991) criteria, these characteristics, combined with the fact that
the species of H. brachyacantha complex are more widely distributed elsewhere (Sun et al.,
2016), support a likely introduced status for the H. brachyacantha complex in the
Mediterranean Sea.
The new records of this complex of species presented from this study in Greece and
Spain demonstrate its ongoing spread, and additionally provide an important reference

for future genetic analyses. Due to the uncertainty surrounding the real identity of any
Mediterranean H. brachyacantha material, we here use the open nomenclature qualifier
“sensu lato.”
Hydroides dirampha Mörch, 1863
Potential native origin: Tropical Western Atlantic.

Distribution: Circumtropical (Bastida-Zavala & ten Hove, 2003), originally described
from the US Virgin Islands (Zibrowius, 1971). It was reported in the Red Sea (Zibrowius,
1971), the Western Atlantic (Bastida-Zavala & ten Hove, 2002), the Eastern Pacific
(Bastida-Zavala & ten Hove, 2003), Australia (Hayes & Sliwa, 2003; Sun et al., 2015), and
Hawaii (Bastida-Zavala, 2008).
In the Mediterranean, it was first reported in Italy in 1870 as Eupomatus lunifer
(Claparède, 1870). It has since spread all over the basin, being next reported in Spain in
1923, Egypt in 1924 (for both records: Zibrowius, 1973), Israel in 1937 (Ben-Eliahu &
ten Hove, 1992), Tunisia in 1969 (Zibrowius, 1978), Lebanon in 1978 (Zibrowius & Bitar,
1981), Turkey in 2005 (Çinar, 2006) and Greece in 2014 (Corsini-Foka et al., 2015).
New records: This finding represents a new country record for Malta (#22 and #23: Figs.
S9D and S9E).
Boat-hull records: Found on boat-hulls moored in marinas in France (#7), Italy (#12,
#15, #17, #20, #21), Malta (#22 and #23), Greece (#24 and #25), Turkey (#31 and #32) and
Cyprus (#33).
Notes: It is a NIS in the Mediterranean believed to be arrived by the shipping pathway
from the tropical Western Atlantic (Zibrowius, 1992).
Hydroides elegans (Haswell, 1883)
Native origin: Australasia and Indian Ocean.

Distribution: Circumtropical: Pacific Ocean, Caribbean, Atlantic and Northern Europe.
In the Mediterranean Sea, it has been reported since the 19th century (Claparède, 1870),
and has since spread to most countries in the basin (Galil & Gevili, 2014).
New records: This finding represents a new country record for Malta (#22). This species
was found in all marinas, except for #8, #13, #14, #20, #23, #29-31, #33, #34, #35. Figs. S9F
and S9G are from #18.
Boat-hull records: Found on boat-hulls from all marinas which had boats sampled.
Notes: It is considered the main fouling organism in the Mediterranean Sea (Koçak, Ergen
& Çinar, 1999); our study confirms that it is the most widespread fouling species found
here in terms of distribution.
Hydroides homoceros Pixell, 1913
Potential native origin: Indo-Pacific.

Distribution: It was originally described from the Cape Verde Islands, in the Eastern
Atlantic (Pixell, 1913). Also reported from the Red Sea, Suez Canal, Arabian Gulf,
Zanzibar and Maldives (Ben-Eliahu & ten Hove, 2011).
Its first Mediterranean record was from Israel in 1955 (Ben-Eliahu, 1991), then in late
1970s from an aircraft carrier moored in Toulon, France (Ben-Eliahu & ten Hove, 2011).
Next it was reported from the south-eastern Turkey (Çinar, 2006).
Boat-hull records: Found on boat-hulls moored in Cyprus (#33: Figs. S9H and S9I), but
was not found in the same marina. The captain of one boat hosting this species in Cyprus
had recently travelled to the Turkish Levantine coast, where it is known from. If it does
establish in Cyprus, it would then present a new country record.
Spirobranchus tetraceros sensu lato (Schmarda 1961)

Distribution: First described from Australia, it has a circumtropical distribution that
includes the Suez Canal, Indian Ocean, South Africa, Australia, Malaysia, Japan, China
and the Caribbean (Ben-Eliahu & ten Hove, 1992; Fiege & Sun, 1999).
Its first Mediterranean record was from Lebanon in 1965 (Laubier, 1966) as
Spirobranchus giganteus coutierei Gravier, 1908 (which is now understood as a sub-species
of Spirobranchus tetraceros; E. Kupriyanova, 2017, personal communication), followed by
Rhodes, Greece (Dumont & Werger, 1989), Abu Kir Bay, the Egyptian Mediterranean
(Selim et al., 2005), and the Turkish Levantine Sea (Çinar et al., 2006).
New records: This finding presents a new subregional record for the Central
Mediterranean and a new country record for Italy (#18: Fig. S9J). It was also found in
Greece (#24).
Notes: Spirobranchus tetraceros has been treated as a complex of species since 1994 (Fiege
& Sun, 1999; Ben-Eliahu & ten Hove, 2011) in need of taxonomic revision, hence, here it is
referred to as Spirobranchus tetraceros sensu lato.
Porifera
Family: Amphoriscidae
Paraleucilla magna Klautau, Monteiro & Borojevic, 2004
Potential native origin: Indo-Pacific and Australia.

Distribution: First described from the Western Atlantic in Rio de Janeiro, Brazil, and was
also found from the Azores, Madeira and Portugal (Bertolino et al., 2014; Guardiola,
Frotscher & Uriz, 2016).
Its first Mediterranean records were from multiple Italian localities, first in the Ionian
then in the Tyrrhenian and Adriatic Seas, followed by the Ligurian Sea and Sicily (Longo
et al., 2004, 2012; Longo, Mastrototaro & Corriero, 2007; Bertolino et al., 2014; Marra et al.,
2016). It was also reported from multiple localites in the Costa Brava region in Spain
(Guardiola, Frotscher & Uriz, 2012, 2016), as well as Malta (Zammit, Longo & Schembri,
2009), and Croatia (Cvitkovic et al., 2013). Its first Eastern Mediterranean record is from

the Gulf of Thessaloniki, Greece, where it was first observed in 2014 in a mussel farm
(Gerovasileiou et al., 2017). It has also emerged in the Sea of Marmara, Turkey in 2012
(Topaloğlu, Evcen & Çınar, 2016).
New records: This finding represents a new country record for Cyprus (#34), and new
locality records for both Greece (#24: Fig. S10A, and #26) and Sicily. Specifically, it was
present at all seven sampled Sicilian marinas (#15–21), and was also found in Malta (#23).
Boat-hull records: Found on boat-hulls in France (#5), Malta (#22), Greece (#26) and
Cyprus (#34).
Notes: Prior to the recent 2004 description of Paraleucilla magna, this genus was only
known from the Indo-Pacific region and Red Sea. As it was initially described from
Rio de Janeiro in 2004, where it is considered cryptogenic (Cavalcanti et al., 2013), it was
likely already present in several Mediterranean locations. For instance, the species had
already been recorded (preceding its formal description) in 2001 from Mar Piccolo of
Taranto (Longo et al., 2004), and according to local mussel farmers was present there as
long as 20–30 years earlier (Longo, Mastrototaro & Corriero, 2007). The opportunistic
behavior of Paraleucilla magna, with proliferation only close to either aquaculture
facilities or harbours, may be the reason behind its late detection in the Mediterranean
(Guardiola, Frotscher & Uriz, 2012). Moreover, as several introductions probably occurred
in a short period of time, the phylogeographic signal could be weak or even lost, making
the determination of the introduction pathway a challenge (Pineda, López-Legentil &
Turon, 2011). Aquaculture and shipping are the most probable vectors for its recent
expansion along the Western Mediterranean coast (Longo, Mastrototaro & Corriero, 2007).
This study shows that Paraleucilla magna is now both a common and established species
around Sicily and Malta. Noteworthy is the record on boat-hulls reported here from
France (but not in the marina), which may represent the first step of subsequent spreading
in the Western Mediterranean and recreational boating as another vector of spread.
Pycnogonida
Family: Ammotheidae
Achelia sawayai sensu lato Marcus, 1940
Native origin: Western Atlantic.

Distribution: Extremely common in the tropical shallow waters of the Western Atlantic. It
is distributed from Georgia to the Gulf of Mexico and throughout the Caribbean Sea to
Brazil (Müller & Krapp 2009). It has also been reported in Western Africa, Madagascar and
in the southern Pacific in French Polynesia, Indonesia, Fiji and Papua New Guinea,
although some of these records are still awaiting confirmation (Child, 1992, 2004).
New records: This finding represents a new record for the Mediterranean Sea, and new
country records for both Malta (#23: Figs. S11A and S11B) and Italy (#17, #18).
Notes: Recent molecular studies suggest that the Atlantic and Pacific populations may
belong to different entities within a complex of species (Sabroux et al., 2017). Therefore, it
is referred to here as Achelia sawayai sensu lato. Since the origin of the Mediterranean

material is unknown, further molecular studies are necessary to understand the invasion
route taken by this pycnogonid. The local reproductive success of this species exhibiting
paternal care was demonstrated by the finding of two ovigerous male specimens.
DISCUSSION
This wide-scale study spanning the Mediterranean Sea provides a massive update of new
NIS records, and in many cases their regional or local expansions, providing a warning for
subsequent spreading. The 51 new country records presented in this study clearly indicate
how inadequate our knowledge on Mediterranean marine NIS still is. There was a
prevalence of new findings for bryozoans and crustaceans in almost all countries (Fig. 2).
This is because these are both poorly studied taxa in the Mediterranean owing to a lack of
taxonomic expertise/focused studies. Additionally, typical rapid assessment surveys or
citizen science initiatives searching for NIS usually target larger and eye-catching taxa
(Zenetos et al., 2013; Mannino et al., 2017), so these minuscule or less charismatic
components of fouling biota may have gone previously overlooked or not have had the
applicable expertise available.
It is not uncommon for marine NIS to go overlooked for long periods of time
(Carlton, 2009), as in the case of Paraleucilla magna in the Mediterranean
(Longo, Mastrototaro & Corriero, 2007). Actually, for most “first country records”
documented in this study, the year of first introduction may have been much earlier than
the first year of discovery presented here, as these NIS may have gone unnoticed due to a
lack of taxonomic expertise or lack of focused study. Some probable examples of this
include: Paranthura japonica, Watersipora arcuata and Celleporaria brunnea, whose
current widespread Mediterranean distributions indicate they have likely been hitching
rides around the basin for quite some time. Another example of this is the sea spider
Achelia sawayai sensu lato, first reported here for the Mediterranean basin, and specimens
were already found in three marinas: two in Sicily and one in Malta. An exception to this is
our finding of Microcosmus exasperatus in Karpaz Gate Marina, Cyprus, as this species was
specifically sought two years prior to our sampling of the same marina, but was not found
then (Gewing et al., 2016). Also, Celleporaria brunnea was not found to be present in
Grand-Motte, France in 2014 (Marchini, Ferrario & Minchin, 2015), but was present
there when we sampled in 2016.
In addition to the records presented here, Percnon gibbesi (H. Milne Edwards, 1853)
was sighted in Port Vauban, France, presenting a new country record for this species of
western Atlantic origin, and already known from several other Mediterranean countries as
a very successful invader (Katsanevakis et al., 2010, 2011). The non-indigenous status of
Percnon gibbesi in the Mediterranean Sea is uncertain, because its long-lived planktonic
larvae could have entered the Gibraltar Strait facilitated by natural means, i.e. the Atlantic
Current, rather than human vectors, such as ballast water (Mannino et al., 2017 and
references therein). Due to its questionable status regarding its mode of introduction, we
have cautiously separated this species from the other NIS records. However, it is
noteworthy that Percnon gibbesi was sighted feasting on a fouling community on a

boat-hull in Greece (#25), suggesting hull-fouling as another possible vector for its
ongoing spread.
This study focusing exclusively on marina habitats indicates that recreational
boating represents the most plausible vector of introductions for the NIS we found, aside
from the few marinas situated in very close proximity to either aquaculture facilities or
shipping ports, as in Port Camargue, France and Heraklion, Greece. Hence, most of these
new records suggest the pivotal role of recreational boating in facilitating both first
introduction events to a given country and secondary spread.
Furthermore, some species reported here are likely polyvectic, but it is clear that
recreational boating plays a determinant role in accelerating/facilitating the spread of
many species, especially those having only a very short and lecithotrophic larval stage.
The presence of such species lacking the ability for natural long-distance dispersal found
on boat-hulls and in marinas can confirm that the hull-fouling vector is instrumental in
expediting primary introductions as well as facilitating secondary transfer for many
ascidians, bryozoans and peracarids such as Ampithoe bizseli, Bemlos leptocheirus,
Celleporaria brunnea, Clavelina oblonga, Paraleucilla magna, Paracerceis sculpta,
Paranthura japonica, Phallusia nigra, Styela plicata and Tricelleria inopinata. The ongoing
nature of the invasion process is further demonstrated by the observation of the same set
of NIS on boat-hulls and in the same marinas, clearly showing the exchange of organisms
from marina to mobile habitats and vice versa.
The species which are not yet present in a country, but found only on boats obviously
cannot formally be recorded as new country records, unless we are certain that the boat
has not left that country since its last hull-painting/cleaning, e.g., as in our finding of
Paradella dianae on boat-hulls in Greece. Some other interesting cases of NIS found on
boats but not yet in the country are (see Table 3 for details) the barnacle Amphibalanus
improvisus and the bryozoan Watersipora arcuata both found on hulls in France, yet the
boats which they were found on had only travelled to the Balearic Islands, alluding to the
assumption that those NIS are likely present in the Balearic Islands. Also noteworthy is the
finding of the oyster Saccostrea glomerata from a boat-hull in France, representing a new
record for the Western Mediterranean, and of the bryozoan Parasmittina egyptica from a
boat-hull in Italy, representing the first Central Mediterranean record for this species.
Overall, the 20 records presented in this study of NIS attached to boats but not yet
recorded in the respective marina illustrates the potential of the biofouling vector in
seeding a new area with propagules.
In synthesis, a pool of NIS is circulating among Mediterranean marinas, linked by a
dense network of boat voyages ensuring their dissemination by a steady multiplication of
the number of occasions. It is also of interest to point out that nearly all marinas have a
rule prohibiting the in-water cleaning of vessels, but this rule is genuinely not enforced,
and in-water cleaning was commonly witnessed within marinas during this study, likely
facilitating the ‘stepping stone’ invasion process by dislodging and exacerbating the
resettlement of NIS propagules.
Recently, Ferrario et al. (2017) showed that marinas can host as many NIS as larger
commercial harbors. This massive contribution of new NIS records confirms their result

and reveals that Mediterranean marinas so far have been inadequately explored for NIS,
despite the Mediterranean Sea being both the global hotspot for boating traffic, and for
level of NIS invasions. We strongly recommend that major attention should soon be
dedicated to recreational marinas as hotspots of introduction, and to pleasure boats as a
vector of introduction and spreading. Management actions to combat NIS in the
Mediterranean Sea need to also incorporate the recreational boating vector.
DEFINITIONS
Non-indigenous species An organism introduced outside its natural past or present
distribution range by human agency, either directly or
indirectly (European Environment Agency, 2012).
Polyvectic species May have been introduced by a certain combination of
vectors and pathways (Carlton & Ruiz, 2005).
Pseudoindigenous species A species described as new to an area where it was in fact
introduced by human action (Carlton, 2009).
ACKNOWLEDGEMENTS
We thank all the marina personnel for granting permission for this study; please see
Supplementary Material for a detailed list of those who facilitated this study. We are also
incredibly indebted to the boat owners/captains for their cooperation. For helping to
obtain marina permissions we thank Giulio Franzitta, Paola Gianguzza and Stefanos
Kalogirou. For collection of samples, we thank Alessandro Bolis, Gina Donnarumma, Ali
Murat Elagoz, Andres Izquierdo-Muñoz, Alberto Orengo, Francesco Saracino, the late
Arif Sipahi and Tevfik Yilmaz. For taxonomic advice, we would like to thank (in
alphabetical order) Michel Bariche, Ferdinando Boero, Şermin Açik Çinar, Melih Ertan
Çinar, Victoria Fernandez-Gonzalez, Bella S. Galil, Jean-Georges Harmelin, Valiallah
Khalaji- Pirbalouty, Traudl Krapp-Schickel, Elena Kupriyanova, Gretchen Lambert, Carlos
Navarro-Barranco, Gary Poore, Rosanna Rocha, Jean-Claude Sorbe, Yanan Sun and Kevin
J. Tilbrook. We are also grateful to Francis Kerckhof and Giuseppe de Paolis for their help
in obtaining copies of publications of difficult access.
ADDITIONAL INFORMATION AND DECLARATIONS
Funding
This work was funded by a PhD Scholarship awarded to Aylin Ulman from the MARES-
Erasmus Mundus Joint Doctoral Fellowship Program in Marine Ecosystem Health and
Conservation. A COST Action #1209 grant was provided to Aylin Ulman to facilitate
ascidian taxonomic identification in the University of Alicante, Spain. MARES is a Joint
Doctorate programme selected under Erasmus Mundus coordinated by Ghent University
(FPA 2011-0016). The funders had no role in study design, data collection and analysis,
decision to publish, or preparation of the manuscript.

Grant Disclosures
The following grant information was disclosed by the authors:
Aylin Ulman from the MARES-Erasmus Mundus Joint Doctoral Fellowship Program in
Marine Ecosystem Health and Conservation.
Ghent University: FPA 2011-0016.
Competing Interests
The authors declare that there are no competing interests.
Author Contributions
Aylin Ulman conceived and designed the experiments, performed the experiments,
analyzed the data, contributed reagents/materials/analysis tools, wrote the paper,
prepared figures and/or tables, reviewed drafts of the paper.
Jasmine Ferrario performed the experiments, analyzed the data, contributed reagents/
materials/analysis tools, wrote the paper, prepared figures and/or tables, reviewed drafts
of the paper.
Anna Occhpinti-Ambrogi conceived and designed the experiments, contributed
reagents/materials/analysis tools, reviewed drafts of the paper.
Christos Arvanitidis performed the experiments, analyzed the data, contributed
reagents/materials/analysis tools, reviewed drafts of the paper.
Ada Bandi performed the experiments, analyzed the data.
Marco Bertolino analyzed the data, contributed reagents/materials/analysis tools, wrote
the paper, prepared figures and/or tables, reviewed drafts of the paper.
Cesare Bogi analyzed the data, contributed reagents/materials/analysis tools, wrote the
paper, prepared figures and/or tables, reviewed drafts of the paper.
Giorgos Chatzigeorgiou analyzed the data, contributed reagents/materials/analysis
tools, reviewed drafts of the paper.
Burak Ali Çiçek contributed reagents/materials/analysis tools, reviewed drafts of the
paper, helped with logistics and permissions.
Alan Deidun contributed reagents/materials/analysis tools, reviewed drafts of the paper,
helped with logistics and permissions.
Alfonso Ramos-Esplà analyzed the data, contributed reagents/materials/analysis tools,
wrote the paper, reviewed drafts of the paper.
Cengiz Koçak analyzed the data, contributed reagents/materials/analysis tools, wrote the
paper, prepared figures and/or tables, reviewed drafts of the paper.
Maurizio Lorenti analyzed the data, contributed reagents/materials/analysis tools, wrote
the paper, reviewed drafts of the paper.
Gemma Martinez-Laiz performed the experiments, analyzed the data, prepared figures
and/or tables, reviewed drafts of the paper.
Guenda Merlo performed the experiments, analyzed the data.
Elisa Princisgh performed the experiments, analyzed the data.
Giovanni Scribano performed the experiments, analyzed the data, prepared figures
and/or tables.

Agnese Marchini conceived and designed the experiments, performed the experiments,
analyzed the data, contributed reagents/materials/analysis tools, wrote the paper,
prepared figures and/or tables, reviewed drafts of the paper.
Field Study Permissions

The following information was supplied relating to field study approvals (i.e., approving
body and any reference numbers):
The following people (including their position) authorised the field sampling
permissions for this study:
Spain
Marina de Alicante: Seaman chief—Marı̀n
One Ocean Port Vell Barcelona: Communications Officer—Regina Sorreno
France
Cap d’Agde: Marina Director—Pierre Weiss
La Grand-Motte: Marina Director—Eric Pallier
Port Camargue: Marina Director—Michele Cavalais
Saint-Tropez: Marina Director—Jean-Francois Tourret
Marines du Cogolin: Marina Director—Claude Robert
Saint-Maxime: Marina Manager—Mr. Ellul
Cannes Le Vieux Port: Marina Director—Francois Gillet
Antibes Port Vauban: Marina manager—Emilie Pluenet
Villefranche-sur-Mer: Chef d’Exploitation—Sarah Castanie
Italy
Porto Turistico di Roma/Lido di Ostia: Marina Manager—Daniele Cossu
Ischia Marina di Sant’Angelo: Giulio Lauro; Porto d’Ischia: Mr. Di Maio
Sorrento Marina Piccola: Italian Coast Guard/Sorrento Location
Marina Villa Igiea Palermo: Marina Owner—Gioacchino Guccione
Porto La Cala Palermo: Marina Manager—Nicola Rossa
Porto Grande Siracusa: The Italian Coast Guard/Siracusa granted verbal permission.
Marina di Marzamemi: Marina Director—Salvatore Gurrieri
Marina di Ragusa: Marina Manager—Enza Di Ramondo
Marina di Cale del Sole (Licata): Marina Manager—Maria Sitibondo
Porto dell’Etna/Marina di Riposto: Marina Manager—Emiliano Indelicato
Malta
Msida Yacht Marina: Marina Director—George Mercieca
Grand Harbour Marina Valletta: Marina Director—Gordon Vassolo
Greece
Old Venetian Harbour (Heraklion): Marina Director—Dr. Bras Ioannis
Agios Nikolaos Marina: Marina Manager—Mikhalis Farsaris and Marina
Director—Garefallakis Michalis.
Rhodes Mandraki Port: Chief of the Port Police—Mr Moustakopoulos Ioannis

Turkey
Istanbul Atakoy Marina: Marina Manager—Asli Ebru Erkoc
Istanbul Setur Kalamiş Marina: Marina Manager—Kerem Cesmebasi
Milta Bodrum Marina: Marina Communications Director—Ayşe Mine Aykutluğ
Setur Marmaris Netsel Marina: Marina Director—Erkan Ozatag & Marina
manager—Onur Kunduz
Datça Marina: Marina Manager—Ali Gök
Ece Fethiye Marina: Marina communications officer—Mrs. Yelena
Setur Finike Marina: Marina Manager—Zia Dal
Cyprus
Karpaz Gate Marina: Marina Director—Deniz Akaltan
Famagusta Port: The Turkish Coast Guard granted verbal permission with Erol
Adelier’s and Burak Çiçek’s assistance.
Data Availability
The following information was supplied regarding data availability:
The data that this output produced is all detailed in Tables 2, 3 and 5, with details of each
species in the Results.
Supplemental Information
Supplemental information for this article can be found online at http://dx.doi.org/
10.7717/peerj.3954#supplemental-information.
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A massive update of non-indigenous

species records in Mediterranean marinas

Subjects Biodiversity, Conservation Biology, Ecology, Marine Biology, Taxonomy

Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 2/59

Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 3/59

MATERIALS AND METHODS

Marina substrate sampling

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Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 5/59

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Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 8/59

Clavelina oblonga Turkey (#29) *

Phallusia nigra Cyprus (#33, #34) *

Styela plicata Malta (#22, #23) *

Symplegma brakenhielmi Italy (#15) *, CM

Amathia verticillata Cyprus (#34) *

Amathia verticillata Turkey (#28, #30) *

Celleporaria brunnea Spain (#1) *

Celleporaria brunnea France (#4, #5, #6, #8) *

Celleporaria brunnea Malta (#22, #23) *

Celleporaria brunnea Greece (#24) *

Celleporaria vermiformis Greece (#24, #25, #26) *

Celleporaria vermiformis Cyprus (#33, #34) *

Hippopodina sp. A Turkey (#32) *

Parasmittina egyptiaca Turkey (#32) *

Parasmittina egyptiaca Cyprus (#33) *

Tricellaria inopinata France (#3, #5) *

Tricellaria inopinata Greece (#24) *

Watersipora arcuata Spain (#1, #2) *

Watersipora arcuata Malta (#22) *, CM

Aorides longimerus France (#5) **

Bemlos leptocheirus Greece (#24, #25) *, EM

Cymodoce cf. fuscina Greece (#24) **

Dyspanopeus sayi Greece (#24) *, EM

Ianiropsis serricaudis France (#3, #5) *

Mesanthura cf. romulea Spain (#1) *

Mesanthura cf. romulea Malta (#22) *

Mesanthura cf. romulea Greece (#26) *

Mesanthura cf. romulea Cyprus (#33, #34) *

Paracerceis sculpta Malta (#22, #23) *

Paracerceis sculpta Cyprus (#34) *

Paranthura japonica Spain (#1, #2) *

Paranthura japonica Malta (#23) *

Sphaeroma walkeri Greece (#24) *

Stenothoe georgiana France (#5) *

Stenothoe georgiana Malta (#23) *, CM

Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 9/59

Dendostrea folium s.l. Malta (#22, #23) *, CM

Hydroides dirampha Malta (#22, #23) *

Hydroides elegans Malta (#22) *

Spirobranchus tetraceros s.l. Italy (#18) *, CM

Achelia sawayai s.l. Malta (#23) **

Pycnogonida Achelia sawayai s.l. Italy (#17, #18) **

NEW NIS RECORDS: NOTES ON INDIVIDUAL SPECIES

Ulman et al. (2017), PeerJ, DOI 10.7717/peerj.3954 10/59

Tricellaria inopinata Turkey (#27) *

Parasmittina egyptiaca Italy (#21) *, CM

Watersipora arcuata France (#7) *

Crustacea Amphibalanus improvisus France (#5) *

Balanus trigonus Cyprus (#33) *

Cymodoce cf. fuscina Greece (#25) L

Paradella dianae Italy (#20) L